Skip to main content

Full text of "The Nautilus"

See other formats 


americanmalacologists,  inc. 

PUBLISHERS  OF  DISTINCTIVE  BOOKS  ON  MOLLUSKS 

THE  NAUTILUS  (Quarterly) 

MONOGRAPHS  OF  MARINE  MOLLUSCA 

STANDARD  CATALOG  OF  SHELLS 

INDEXES  TO  THE  NAUTILUS 

{Geographical,  vols  1-90;  Scientific  Names,  vols  61-90) 

REGISTER  OF  AMERICAN  MALACOLOGISTS 


JANUARY  31,  1986 


THE 


NAUTILUS 


FEB    5  1986 

I     Woods  Hole,  Mass. 


ISSN  0028-1344 


Vol.  100 


No.  1 


A  quarterly 

devoted  to 

malacology  and 

the  interests  of 

conchologists 


Founded  1889  by  Henry  A.  Pilsbry.  Continued  by  H.  Burrington  Baker. 
Editor-in-Chief:  R.  Tucker  Abbott 


EDITORIAL  COMMITTEE 
CONSULTING  EDITORS 


Dr.  William  K.  Emerson 
Department  of  Living  Invertebrates 
The  American  Museum  of  Natural  History 
New  York,  NY  10024 

Mr.  Samuel  L.  B.  Fuller 
1053  Mapleton  Avenue 
Suffield,  CT  06078 

Dr.  M.  G.  Harasewych 
363  Crescendo  Way 
Silver  Spring,  MD  20901 

Mr.  Richard  I.  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Los  Angeles  County  Museum  of  Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 


Dr.  Arthur  S.  Merrill 
c/o  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Donald  R.  Moore 
Division  of  Marine  Geology 
School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 

Dr.  G.  Alan  Solem 
Department  of  Invertebrates 
Field  Museum  of  Natural  History 
Chicago,  IL  60605 

Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Gilbert  L.  Voss 
Division  of  Biology 

School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 


EDITOR-IN-CHIEF 

Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
Box  2255,  Melbourne,  FL  32902-2255 

Mrs.  Cecelia  W.  Abbott 
Business  and  Subscription  Manager 
P.O.  Box  2255 
Melbourne,  FL  32902-2255 


Second  Class  Postage  paid  at  Melbourne,  Florida 
and  other  post  offices 


The  Nautilus  (USPS  374-980) 
ISSN  0028-1344 

A  quarterly  magazine  devoted  to  malacology. 
Copyright  ^1986  by  American  Malacologists,  Inc. 

OFFICE  OF  PUBLICATION 

American  Malacologists,  Inc.  (United  Parcel  Address: 

2208  South  Colonial  Drive,  Melbourne,  FL  32901) 

Mail:  Box  2255,  Melbourne,  FL  32902-2255 

POSTMASTER:  Send  address  changes  to  above. 

Subscription  Price:  $15.00  (see  inside  back  cover) 

$17.00  (foreign);  institutions  $20.00 


THE 

NAUTILUS 

Volume  100,  number  1  —  January  31,  1986 

ISSN  0028-1344 


CONTENTS 

Dedication  of  the  100th  Volume 1 

Birth  and  Growth  of  The  Nautilus 2 

Tom  Pullev  and  the  Train 8 


Joseph  Rosewater  (1928-1985) 

A  Tribute  and  Bibliography  by  Harald  A.  Rehder . 


Andrew  C.  Miller,  Barry  S.  Payne  and  Terry  Siemsen 

Description  of  the  Habitat  of  the  Endangered  Mussel  Plethobasus  cooperianus 14 

Andrew  C.  Miller,  Barry  S.  Payne  and  David  W.  Aldridge 

Characterization  of  a  Bivalve  Community  in  the  Tangipahoa  River,  Mississippi 18 

Richard  A.  Petit 

Notes  on  Species  of  Brocchinia  (Gastropoda:  Cancellariidae) 23 

William  K.  Emerson 

On  the  Type  Species  oi Metula  H.  &  A.  Adams,  1853: 

Buccinum.  clathratum  A.  Adams  and  Reeve,  1850  (Gastropoda:  Buccinidae) 27 

Eugene  Coan 

Some  Additional  Taxonomic  Units  that  First  Appear  in  Publications  by  J.  G.  Cooper 30 

Dee  S.  Dundee 

Notes  on  the  Habits  and  Anatomy  of  the  Introduced  Land  Snails, 

Rumina  and  Lamellaxis  (Subulinidae) 32 

Award  News 37       Notes;  Deaths 38 


STATEMENT  OF  OWNERSHIP,  MANAGEMENT  AND  CIRCULA- 
TION (Required  by)  Act  of  October  23,  1962:  Section  4396,  Title 
39.  United  States  Code,  and  postal  regulation  132-622. 

1.  Title  of  publication    THE  NAUTILUS. 

2.  Date  of  filing:   September  14,  1985 

3.  Frequency  of  Issue:   Quarterly  (4  per  year). 

4.  Location  of  known  office  of  publication:  2208  South 
Colonial  Dr.,  Melbourne,  FL  32901. 

5.  Location  of  Headquarters  of  General  Business  Offices 
of  the  Publishers:  2208  South  Colonial  Dr.,  Melbourne,  FL 
32901 , 

6.  Names  and  addresses  of  publisher,  editor,  and  manag- 
ing editor:  Publisher,  American  Molocologists,  Inc.,  P.O. 
Box  2255,  Melbourne,  FL  32901,  Editor,  R.  Tucker  Abbott, 
P.O.  Box  2255,  Melbourne,  FL  32902,  Business  Manager, 
Mrs.  Cecelia  W  Abbott,  P.O,  Box  2255,  Melbourne,  FL 
32902-2255, 

7.  Owner:  American  Molocologists,  Inc.,  P.O.  Box  2255, 
Melbourne,  FL  32902-2255. 

8.  Known  bondholders,  mortgages,  and  other  security 
holders   owning   or    holding    1    percent   or    more   of   total 


amount  of  bonds,  morfoges  or  other  securities:    none. 
9,    Extend  and  Nature  of  Circulation: 


Average   Single 

12  Mos.      Issue 

850  850 


A.  Total  No.  copies  Printed  (Net  Press) 

B.  Paid  Circulation 

1.  Soles  through  dealers  and  carriers, 
street  vendors  and  counter  sales 

2.  Moil  subscriptions 

C.  Total  Paid  Circulation 

D.  Free  Distribution  (including  samples) 
by  moil  corner  or  other  means 

E.  Total  Distribution  (Sum  of  C&D) 

F.  Office  use,  left-over,  unaccounted 
and  back  start  subscription  copies 

G.  Total  (Sum  of  ESF)-should  equal  net 
press  run  shown  in  A. 


I  certify  that  the  statements  mode  by  me  above  are  correct  and 

complete. 

(signed)  R.  Tucker  Abbott,  Editor 


none 

none 

735 

730 

735 

730 

18 

18 

753 

748 

91 

102 

850 

850 

THE  BEST  OF  THE 


An  Exciting  and  Nostalgic  Glimpse 
Into  Conchologys  ftist 


Heii'  IS  a  IreiTSLirv  of  iinsl.Tlf^n  and  ofton 
humorous  writings  takfii  from  tho  first  40 
voars  of  America's  oldest  mollusk  |ourn,il, 
covering  the  Lite  pioneering  f^'riod  of 
1886  to  1927  The  noted  m.iKicoiogist  and 
editor,  R  Tucker  Abholl  has  carefully 
gathered  a  wide-ranging  series  of  over 
100  articles  that  depict  tiir  lore  and  ex- 
citement of  sli.'il  hunlmt;  in  the  United 
States 


American  M«lacologlst5 

P.O.  Bo.  2255  288  pp  ,  rinthbound     $13,95. 

Melbourne,  FL  32902-2255 


Collectible  Shells 

of  Southeastern  U.S.,  Bahamas 

&  Caribbean  by  R,  Tucker  Abbott.  Ph  d 
A  'Take  it  to  ttie  Beach'  Field  Guide 
WATERPROOF  -  TEARPROOF 
105  beautiful  color  photos  of  living  animals  and 
their  shells  64  pages  ol  color  300  species  il- 
lustrated. How  to  clean  shells.  Where  to  find 
them.  Includes  fossils,  pond  and  tree  snails,  as 
well  as  sealife 

Collectible  Shells  stresses  conservation,  but 
also  has  helpful  hints  about  collecting  and 
cleaning  shells.  The  book  introduces  the  tourist 
and  beginner  to  famous  Florida  fossils  and  the 
unique  world  ol  tree  and  pond  mollusks 

Printed  on  a  washable,  tearproof  plastic  paper. 
Drop  it  in  the  ocean,  use  it  in  the  ram,  or  let  your 
wet  shells  drip  all  over  it.  Keep  it  on  your  boat  or 
take  It  to  the  shore  A  popular  new  seller  retailing 
(or  $8.95  Postage  a  id  state  lax  are  included  as  a 
big  savings 


American  Malacologists,  Inc. 

Publishers  of  Distinctive  Books  on  Mollusks 
P.O.  Box  2255,  Melbourne,  FL  32902-2255 

W«  accept  VISA  Of  MASTERCARD  orders  Dymai'  Piea";e  give  dale  o'  e« 
pifation  »n(j  youf  ctfd  number  Fo'eigr*  customers  may  send  mte'na 
lionai  postal  money  order  ot  checK  on  New  Yofh  bank  o'  U  S  cash  by 
'egisiered  mail 


Vol.  100(1)  January  .31,  1986  THE  NAUTILUS 


The  Nautilus 

ITS  100th  ANNIVERSARY 

and 
Volume  100 

Dedicated  to  all  the  mollusk  enthusiasts,  both  private  concholog-ists  and  pro- 
fessional malacologists,  who  founded  and  carried  on  for  the  last  one  hundred 
years  America's  oldest  journal  devoted  exclusively  to  the  study  of  mollusks.  To 
its  500  contributors  and  15  consulting  editors  are  added  many  thanks  and 
appreciation. 

DEDICATED  TO  THE  FORMER  EDITORS^ 

William  D.  Averell  (1853-1928) 
Founder  of  the  Conchologists'  Exchange,  1886 

Henry  A.  Pilsbry  (1862-1957) 
Founder,  editor  of  The  Nautilus,  1888-1957 

Charles  W.  Johnson  (1863-1932) 
Business  manager,  1890-1932 

Horace  Burrington  Baker  (1889-1971) 

Business  manager,  co-editor  1932-1957 

Editor,  1958-1970 

Charles  B.  Wurtz  (1916-1982) 

Co-editor,  1958-1974; 
consulting  editor,  1974-1982 

Bernadine  B.  Baker  (1906-living) 
Business  manager,  1958-1979 

Morris  K.  Jacobson  (1906-1980) 
Consulting  editor,  1972-1980 

William  J.  Clench  (1897-1984) 
Consulting  editor,  1972-1984 

Joseph  Rosewater  (1928-1985) 
Consulting  editor,  1972-1985 


'The  present  staff  is  R.  Tucker  Abbott,  co-editor,  1958-1969, 
editor  since  1970;  Cecelia  W.  Abbott,  business  manager, 
since  1979;  M.  G.  (Jerry)  Harasewych,  associate  editor, 
beginning  with  this  volume.  Mrs.  H.  B.  Baker,  former 
business  manager,  lives  in  Haverford,  Pennsylvania. 


2        THE  NAUTILUS  January  31,  198f; 

BIRTH  AND  GROWTH  OF  THE  NAUTILUS 


Vol.  100(1) 


It  was  the  year  1886.  The  country  was  recov- 
ering from  the  depression  of  1884.  President 
Grover  Cleveland,  in  his  first  term,  was  about  to 
dedicate  the  newly  erected  Statute  of  Liberty  in 
New  York  harbor.  There  were  still  12  states  to 
be  added  to  the  Union,  and  it  took  a  two-cent, 
brown  postage  stamp  with  Washington's  like- 
ness to  send  a  letter  across  the  country.  In 
Philadelphia  the  only  means  of  transportation 
were  horses,  bicycles,  the  steam  locomotive  and 
boats  on  the  Delaware  River.  The  Academy  of 
Natural  Sciences  of  Philadelphia  had  not  built 
its  large  new  annex  and  still  used  gas  lights  in 
its  halls  and  study  rooms.  In  England,  they  were 
celebrating  Queen  Victoria's  Diamond  Jubilee, 
and  the  Spanish  American  War  was  still  12 
years  away. 

In  1886,  in  some  respects,  it  was  the  best  of 


times  for  American  conchology.  William  H.  Dall 
of  the  U.S.  Geological  Survey  in  Washington, 
D.C.  had  already  produced  over  50  major  works 
on  mollusks,  and  Harvard  University  had  just 
published  the  first  part  of  his  famous  "Blake 
Report"  on  deepsea  mollusks.  George  W.  Tryon, 
Jr.,  the  Philadelphia  author  of  "Structural  and 
Systematic  Conchology,"  had  just  completed  the 
tenth  volume  of  his  monumental  "Manual  of 
Conchology."  Verrill  and  Bush  in  New  England 
were  at  the  height  of  their  reports  on  the  dredg- 
ings  of  the  Steamers  FUh  Hnivk  and  the 
AlbatToss.  William  G.  Binney,  living  in  New 
Jersey,  just  across  the  river  from  Philadelphia, 
had  just  revised  his  classic  "Manual  of  North 
American  Land  Shells."  The  Conchological  Sec- 
tion under  the  wing  of  Philadelphia's  Academy 
was  nourishing  with  such  active  members  as  S. 


M 


/  y  y  '    ^ 


7       /  //    y 


o-vSr  ^-^  '    -'^/^ 


■J  11  ^Hy//X 


Tho  end  (if  a  letter  written  in  Dec.  18i»0  to  the  editors  of  the  newly  launc-hiMl  XnuiWux  hy 
the  Calil'ornia-liased  shell  collector  and  newspaper  editor,  Kdward  Warren  Ro(ier,  admon- 
ishing them  to  have  it  "come  out  on  time." 


Vol.  100(1)  January  31,  1986  THE  NAUTILUS        3 

$1.00  per  Year.       ($1.12  to  Foreign  Countries.)       lOcts.  a  copy. 

THE 

NAUTILUS 

A  MONTHLY 

DEVOTED  TO  THE  INTERESTS 

OF  CONCHOLOGISTS. 

EDITOR: 
H.  A.  PiLSBRT,  Conservator  Conchological  Section,  Academy  of  Natural  Sciences,  Philadelphia. 

ASSOCIATE  EDITOR  : 
C.  W.  Johnson,  Acting  Curator  Wagner  Institute  of  Science. 


Vol.  VI.  FEBRUARY,    1893.  No.  10. 

CONTENTS: 

PAGE. 

Additional  Shells  from  the  Coast  of  Southern  Brazil.     Wm.  H. 

Dall 109 

Description  OF  A  NEW  FORM  OF  CYPR.iiA.     John  Ford 112 

Vol.  84 JULY,  1970 No.  1 

THF 

NAUTILUS 

THE  PILSBRY  QUARTERLY 
DEVOTED  TO  THE  INTERESTS  OF  CONCHOLOGISTS 

EDITORS  AND  PUBLISHERS 

R.  Tucker  Abbott,  du  Pont  Chair  of  Malacology 
Delaware  Museum  of  Natural   History.  Greenville.   Del.   19807 

Horace  Burrington  Baker,  11  Chelten  Road,  Havertown,  I'a. 
(Emeritus  Professor  of  Zoology,  University  of  Pennsylvania) 

Charles  B.  Wurtz,  Biology  Department 
La  Salle  College,  Philadelphia,  Pa.  19141 


CONTENTS 

The  occurence  of  Cymatiidae  and  Cypraeidae  in 

North  Carolina.      By  Hugh  J.  Porter 1 

Eggs  and  attachment  sites  for  egg  capsules  of  Valvata  lewisi. 
By  B.  Z.  Lang  and  N.  O.  Dronen,  Jr 9 


THE  NAUTILUS 


January  81,  1986 


Vol.  100(1) 


Vol.  I.  No.  1. 

This  is  the  first  issue  of  "The  Conchologist's 
Exchange  "  As  encouragpnient  is  received  it  will 
assume  the  form  of  a  printed  sheet  with  columns  for 
"  Exch.-inges  in  Mollusca,"  "New  Localities,"  "Answers 
to  Correspondents,"  ike.  This,  our  first  number,  has 
been  sent  to  500  Conchologists.  Subscription  price,  25 
cents  per  annum,  post  paid.  Exchanges  of  20  words,  10 
cents;  for  each  additional  10  words  the  charge  will  be  5 
cents.  The  Conchologist's  Exchange  will  be  issued 
semi-monthly,  and  will  endeavour  to  become  a  cheap 
and  useful  medium  for  the  exchange  of  those  most  beauti- 
itful  productions  of  nature — "The  Mollusks." 

EXCHANGES   FOR   MOLLUSCA  ONLY. 


CVPR.^A  erosa,  L.  lynx,  L. 
CERITHIUM.     maculosum, 

Kien.     eburncum  Brug. 
CVCLOS  rOMA  sulcatum.  Lam. 

clegans  Mull, 
LVMN.-EA  lebraTryon 
bTROMBINA  ^icanahfcra  Sby. 

Fi-^urclla    volcano.    Rve.    Col- 

umbella  fulgunns  Lam. 
Prof.  D.  S.  SHELDON. 

Davenport,  fa 

SUCCINEApuiris  L. 
HLLIX  arbuslorum  L. 
"       ncmomlii  L 
"       ericelorum.  Mull. 

Toiundato,  MuU. 
"       Upicida  L  ccllaria  Mull 
Pupa  muscorum,  L. 
Cionella  subcylindriea 

E    LtHNERT, 
Washington,  D  C. 


CONIOBASIS  simplex,  Suy. 
cariiiifcra.  Lam.  bella,  Con. 
p«rangulata.  Con, 
sordiJa  L«a   symiTielrica,  Hald 
;bcnum.  Lea 

MeUntbo  subsolida  Anth. 
Unioriibigjnosus  Lea 
pustulosus  Lea.  gracilis,  Barnes 

w.  a;  marsh. 

Aledo.  Illinois 


HELIX  albolabris.   Say.     alter- 
nata.  Say.  clauaa.Say. 

elevata,  bay.         fallax,  Say. 
hirsuta,  Say.         inflecta,  Say. 
solilaria,  Say.        monodon, 
Rackett.  Sayii.         Binncy. 

Pupa  armifera.  Say,  corlicana. 
Say.  Fallax,  Say  Unio  cicgans 
Lea,  lachrymosus.  Lea,  parvus, 
Barnes 

EDWARD  A.  ENOS, 

Connersvillc,  Indiana. 


NASSAfossataGld. 
Purpura  saxicola  Val. 
Amycla  gausapata  Gas. 
Adula  falcata  Gld. 
Acmaea  spectrum  Esch 
scabra.  Nuti,  pelta,  Esch. 
Hipponyx  cranioidcs.  Carp. 
G.  W.  PUIFRBAL'GH, 

Greenfield,  Indiana. 


American     and     Foreign 
Unionidac  for  exchange. 
Send  for  list. 


No  responsibility  will  be  ai- 
umed  for  the  siaQ(Jing  of  the 
bovc  parties. 


Address,  WILLIAM  0.  AVERELL,  Proprietor, 

CHESTNUT  HILL,  PHILADELPHIA. 


f  ^e  Conr^ologists'  ^H^ange. 


A  Monthly  Publloatlon  desltrned  for  Ood  .holOitlerta  and  Sclentlsta 
arenerally.     "Wm,  D   Averell.  Editor  and  Publlaher. 


Vol.  II- 


MARCH  and   APRIL,  1888. 


No.  9. 


oojsra?E2sra?s. 


Ill,,  Wm   A. 
H,  Wnght, 


Tbc  .Sl.ell-Hcwing  MuUuscb  of  Kliode  Island,  II,  K,  C»ipenier, 

brief  Noics  on  the  LAod  and  Frcsh-Wner  Shells  of  Mercer  Co. 

Note*  on  the  Umonidaeof  Florida,  Dr,  S.  H.  Wnght  and  Berlii 

Lyogyrus.  Gill,  and  oiher  American  shells,  H.  A    Pibbry, 

MarganuiJA  Hildrcthiaaa.  Lea.  U.  Shimek,     ... 

Notes  on  .^mencao  Shells,  Rev.  Wm.  M.  Beauchamp, 

A  Noted  Scienust  Dead  (Geo,  W.  Tryon,  Jr )     From  Phila.  Public  Ledger, 

Bell  Taps. 

Editorial  Note*,  ,,..... 

The  CoDchologiSt  in  Bermuda,  J,  Matthew  JoDe«. 

A  Collecung  Trip  to  On»et  Bay,  Maw,.  F.  C.  Baker,  . 

Exchangei,  ...... 

Suodard  Work*  oa  Conchology 


109 

110 


"5 
116 
"7 
118 
119 
119 
120 


CHESTNUT  HILL,  Philadelphia,  I'a. 


Slnylo  copies,  C  cents;  50  oenia  a  year  In  the  United  States 
and  Canada;  Foreign  Countries,  62  cents;  Countrtee  not  In 
Universal    Postal  Union,  7B  cents. 


Vol.  lUU(l) 


January  31, 1986 


THE  NAUTILUS 


Raymond  Roberts  (Cypraeidae),  John  Ford 
(Olividae),  Dr.  Benjamin  Sharp  (Scaphopoda) 
and  Angelo  Heilprin  (Florida  fossils). 

But  in  the  late  1880's,  in  other  respects,  it  was 
the  worst  of  times  for  American  conchology.  It 
was  like  the  changing  of  the  guard.  Aging  Isaac 
Lea  died  in  1886  at  the  age  of  95;  Lucy  Say, 
widow  of  America's  first  conchologist,  died  on 
Nov.  15,  1887,  and  Andrew  Garrett,  the  con- 
chologist of  the  South  Seas,  died  the  same 
month.  Then,  like  a  clap  of  thunder,  George 
Tryon,  at  the  age  of  49,  succumbed  to  a  heart 
attack  on  February  5,  1888,  soon  after  a  walk  in 
a  severe  winter  storm.  It  was  a  time  when  there 
was  no  journal  in  America  devoted  solely  to  mol- 
lusks.  Tryons  American  Journal  of  Conchology 
had  succumbed  in  the  financial  panic  of  1873. 

In  Europe,  in  1886,  there  were  eight  respecta- 
ble scientific  journals  devoted  solely  to  mollusks 
-two  each  in  England,  Germany  and  France 
and  one  each  in  Belgium  and  Italy  (Jutting  and 
Altena,  1958).  For  their  outlets,  American 
malacologists  depended  on  the  general  publica- 
tions of  the  Smithsonian  Institution,  the  Califor- 
nia Academy  of  Sciences,  Harvard's  Museum  of 
Comparative  Zoology,  and  the  Academy  of 
Natural  Sciences  of  Philadelphia. 

The  need  for  a  new  American  serial  publica- 
tion was  evident.  There  were  several  hundred 
active  conchologists  in  United  States  and 
Canada,  many  of  them  already  publishing 
privately  or  in  obscure  natural  history 
magazines.  Among  them  were  Henry  Hemphill, 
Temple  Prime,  R.  E.  C.  Stearns,  T.H.  Aldrich, 
W.  W.  Calkins,  Charles  T.  Simpson  and  Victor 
Sterki,  to  mention  but  a  few. 

The  Nautilus  had  a  very  inauspicious  begin- 
ning. William  D.  Averell,  a  33-year-old  shell 
dealer  and  close  friend  of  Tryon's  living  in  near- 
by Chestnut  Hill,  began  the  predecessor  of  The 
NautiliLs  as  the  Conchologists'  Exchange  in  July 
1886.  Volume  1,  number  1  was  merely  a  post- 
card sent  out  to  500  people  announcing  that  the 
annual  subscription  price  for  this  monthly 
4-page  publication  would  be  all  of  25  cents.  It 
was  to  contain  conchological  news,  exchange 
notices  and  answers  to  correspondents.  By 
February  1887,  it  had  grown  to  8  pages  and  the 
price  rose  to  35  cents.  The  final  number,  vol.  2, 
no.  9,  issued  in  April  1888,  two  months  after 
Tryon's  death,  had  16  pages.  Already  it  had 


served  as  a  publication  medium  for  such  well- 
known  mollusk  workers  as  F.  C.  Baker,  Josiah 
Keep  of  California,  C.  F.  Ancey  of  Algiers,  W. 
H.  Dall,  T.  D.  A.  Cockerell  and  a  Harry  A. 
Pilsbry  (later  to  call  himself,  Henry). 

But  a  quirk  of  circumstances  in  Philadelphia 
changed  the  history  of  this  journal.  William  B. 
Marshall,  a  Jessup  Student  at  the  Academy  and 
the  assistant  to  Tryon,  left  in  1887  to  be  the 
Assistant  Zoologist  at  the  New  York  State 
Museum  in  Albany,  and  later  to  serve  under  Dall 
and  Bartsch  in  Washington  for  40  years.  Just  at 
this  time  a  young  newspaper  reporter  by  the 
name  of  Pilsbry  from  Davenport,  Iowa,  and  a 
contributor  to  the  Conchologists'  Exchange, 
visited  the  Academy  on  his  way  to  seek  new 
employment  in  New  York.  Tryon  was  favorably 
impressed  with  the  24-year-old  amateur  con- 
chologist who  already  had  considerable  know- 
ledge about  mollusks,  was  an  excellent  drafts- 
man and  had  a  natural  bent  for  research  and 
publishing.  Upon  Marshall's  sudden  departure 
for  Albany,  Tryon  immediately  invited  Pilsbry 
to  return  from  New  York  and  be  his  new  assis- 
tant. Pilsbry  arrived  in  November  1887,  and 
Tryon  died  three  months  later. 

Pilsbry  was  elected  as  conservator  of  the  Con- 
chological Section  in  1888,  and  he  immediately 
plunged  into  the  task  of  finishing  the  marine 
series  of  the  Manual  of  Conchology.  a  publication 
that  supplied  much  of  his  salary.  There  was  a 
reawakening  of  interest  in  the  Conchologists' 
Exchange,  and  in  early  1889  a  "Prospectus"  was 
sent  out  announcing  its  continuation  as  The 
Na^dihis  with  Pilsbry  as  editor  and  Averell  as 
business  manager.  Volume  3,  no.  1  began  on 
May  5,  1889. 

According  to  Pilsbry's  colleagues  (H.  B. 
Baker,  1958)  the  one  year's  association  with 
Averell  "was  none  too  happy"  and,  at  the  end  of 
volume  4,  no.  1,  Averell's  share  was  bought  out 
by  another  Philadelphian,  at  the  Wagner  Free 
Institute  of  Science:  Charles  W.  Johnson.  This 
ideal  partnership  lasted  for  the  next  42  years 
until  Johnson's  death  in  Boston  in  1932. 

Johnson  was  a  very  affable  and  knowledgable 
gentleman  whose  forte  was  entomology  and 
paleontology,  as  well  as  malacology.  He  was 
particularly  successful  in  inspiring  young 
naturalists,  and  when  he  moved  to  Boston  in 
1903  to  assume  the  curatorship  of  the  Boston 


THE  NAUTILUS 


January  31,  1986 


Vol.  lOU(l) 


Society  of  Natural  History,  William  J.  Clench  as 
a  youngster  fell  under  his  spell.  Years  later, 
from  1926  to  1932,  Clench  was  the  Curator  of 
Mollusks  at  nearby  Harvard,  and  often  assisted 
Johnson  in  the  business  affairs  of  The  Nautilus. 
It  was  Johnson's  understanding  that  Clench 
would  succeed  him  as  business  manager,  but 
upon  the  former's  death  in  1932,  Pilsbry  (then 
69)  thought  that  both  editors  of  The  Nautilus 
should  be  in  the  same  city;  so  H.  Burrington 
Balcer,  one  of  America's  leading  land  snail  ex- 
perts and  Professor  of  Zoology  at  the  University 
of  Pennsylvania,  began  his  association  as  junior 
editor  and  business  manager  in  October  1932 
with  volume  46.  Meanwhile,  Clench  in  1941 
launched  the  marine  journal,  Johnsonia,  naming 
it  after  his  old  mentor. 

The  Nautilus  continued  to  flourish  and  grow. 
Its  history  is  well-told  in  the  Pilsbry  memorial 
number  of  The  Nautilus  by  Dr.  Baker  (1958,  vol. 
71,  no.  3,  pp.  112-115).  Pilsbry  died  on  October 
26,  1957  at  the  age  of  94,  and  the  editorship  of 
the  journal   continued  under  Baker's  capable 


hands  until  his  death  on  March  11,  1971,  at  the 
age  of  82. 

In  April  1958,  two  junior  editors  joined  The 
Nautilus,  one  being  the  land  snail  expert  and 
aquatic  biologist,  Charles  B.  Wurtz  (1916-1982) 
who  had  received  his  Ph.D.  under  Dr.  Baker  at 
the  University  of  Pennsylvania.  At  the  age  of  34 
I  had  left  the  Smithsonian  Institution  in  1954  to 
become  the  first  occupant  of  the  newly-formed 
Pilsbry  Chair  of  Malacology  at  the  Academy.  I 
became  the  other  junior  editor  and  often 
assisted  Pilsbry  in  the  preparation  of  the  jour- 
nal, sometimes  entirely  assembling  and  editing 
it  during  Pilsbry 's  winter  sojourns  in  Lantana, 
Florida.  Also,  at  this  time.  Dr.  Baker's  wife, 
Bernadine  Barker  Baker,  known  to  her  family 
and  friends  as  "Bunny,"  took  over  the  business 
and  subscription  management,  and  continued  in 
that  capacity  with  great  success  for  the  next 
twenty  years. 

In  1969  I  helped  found  and  organize  the 
Delaware  Museum  of  Natural  History  in  Green- 
ville, and  became  the  curator  of  their  Depart- 


fi^osfEcyds,- 


"We  propose  lo  Issue  a  sjxieen  paQ,e  iliustraied 
acTavQ  monrnly  journal  ol  Concholosy  to  be  trailed 

THE   NAUTILUS. 

It  will  lake  ihe  place  cf  ihe  Concholoaist's  Ex- 
change formerly  published  oy  Wm.  D  Averell,  and 
will  be  the  successor  of  that  paper.  Subscribers  lo  the 
Exchange  will  be  crediied  on  the  books  ol  the  Nautilus 
wlih  such  amounts  as  may  be  due  thern  upon  their 
subGCrlptlons. 

The  Nautilus  will  be  under  the  editorial  man- 
agement of  Mr.  Henry  A.  Pilsbry,  Conservator  of  the 
Conchologlcal  Section  at  the  Academy  ot  Natural  Sci- 
ences, and  the  successor  ot  the  late  Mr.  Tryon  in  the 
publication  of  The  Manual  of  Conchology. 

The  new  Journal  is  designed  to  afford  a  medium 
of  communication  between  siudents  and  others  interesi- 
ed  in  conchology  [  to  publish  original  articles  contrib- 
uted by  amencan  and  foreign  authors^  to  record  new 
discoveries)  and  lo  teach  beglnnerG  and  young  studenis 
of  science. 

America  needs  a  Journal  ol  this  kind,  and  if  we 
can  obtain  sufficient  encouragement  In  the  shape  ol 
subscriptions  we  will  commenca  the  publication  of  The 
Nautilus  at  once.  We  enclose  yju  a  blank  for  your 
subscription.  If  you  think  well  of  the  prjject  please 
fill  it  up  at  once  and  return  it  to 

Mouni  Airy,  PriUadalphic.  Pi 


The  Nautilus. 


JANUARY,  1896. 


TO  C0NCH0L0QIST8. 


We  regret  the  necessity  which  compels  ua  to  start  the  new  year 
with  an  editorial  of  this  nature;  but  it  is  nothing  new  to  hear  that 
the  Nautilus  must  struggle  for  existence.  This  struggle  has  con- 
tinued since  the  Jura,  until  now  we  have  but  a  few  species,  three  or 
four  in  the  Indo-Pacific  and  one  in  the  United  States.  Do  vou  in- 
tend that  the  only  recent  Nautill's  In  North  America  shall  be- 
come extinct  ?  We  are  willing  lo  supply  the  care,  but  not  the  en- 
tire environment.  It  needs  feeding  once  a  nionth  in  order  to  add 
another  septum.  You  are  asked  to  assist  in  this  important  function 
once  a  year  ;  and  when  you  see  a  slip  of  paper  which  reads,  "  Inclosed 
please  find  $1,00,"  it  mean.s  that  it  hasconie  your  turn  to  "  chip  in." 
We  hope  that  you  will  no  longer  neglect  these  little  reminders. 
They  mean  that  your  subscription  is  due.  We  cannot  wait  until  the 
end  of  the  year— U'<;  mt(a(  have  \i  in  advance. 

Wishing  you  all  a  Happy  New  Year. 

H.  A.  P.  A  C.  W.  J. 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS 


ment  of  Mollusks.  I  moved  The  Ndutilns.  now 
owned  by  Mrs.  Baker,  to  Delaware.  In  1979, 
upon  the  recommendation  of  Bunny,  my  wife, 
Cecelia  White  Abbott,  took  over  the  duties  of 
business  manager. 

There  was  need  for  an  improvement  in  the 
format  of  the  journal  which  had  remained  un- 
changed for  several  decades.  In  May  1972 
(volume  86)  the  page  size  was  increased  to  8  x 
IOV2  inches,  a  two-column  format  established  on 
a  glossy  paper,  and  a  board  of  13  consulting 
editors  formed.  By  now,  85  years  after  its  incep- 
tion, the  annual  subscription  rate  had  risen  from 
25  cents  to  $7.00  for  individuals.  Today  it  stands 
at  $15.00. 

With  the  demise  of  the  Department  of 
Mollusks  at  the  Delaware  Museum  in  1979  as  a 
functioning,  scientific  entity,  we  moved  The 
Nautilus,  now  under  the  jurisdiction  of  my 
American  Malacologists,  Inc.,  to  Melbourne, 
Florida,  where  it  has  been  published  for  the  last 
six  years.  We  have  now  added  an  associate 
editor,  Dr.  M.  G.  (Jerry)  Harasewych,  a  newly 
appointed  associate  curator  in  mollusks  at  the 
U.S.  National  Museum  in  Washington,  D.C. 

For  over  twenty  years  The  Nautilus  served  as 
the  official  organ  of  the  American  Malacological 
Union  at  no  cost  to  their  organization.  Proceed- 
ings of  the  meetings  were  published  by  them 
from  1932  to  1953  in  the  same  format  as  that  of 
The  Nautilus.  Like  all  growing  organizations, 
the  AMU  eventually  required  its  own  periodical 
and  now  issues  the  extensive  American  Mala- 
cological Bulletin  {vo\.  1,  no.  1,  July  1983)  under 
the  capable  editorship  of  Dr.  Robert  S.  Prezant 
of  the  University  of  Southern  Mississippi. 

The  life  of  an  editor  is  not  easy,  especially  in 
the  case  of  scientific  journals  that  constantly 
need  financial  help.  The  editors  of  the  privately 
owned  Nautilus  were  no  exception,  and  they 


had  to  make  repeated  appeals  for  subscribers  to 
"pay  up."  (Pilsbry  and  Johnson,  1893).  In  my  28 
years  of  experience  as  an  editor  I  had  to  face 
financial  and  production  jiroblems,  and  I  some- 
times found  manuscripts  poorly  written,  badly 
organized,  without  nomenclatorial  knowledge 
and  often  lacking  references  to  the  very  same 
subject.  It  is  work  getting  some  of  them  into 
shape,  and  sometimes  there  is  little  thanks. 
There  is  an  old  adage  that  "the  next  time  you  see 
an  understanding  and  beloved  editor,  you  will 
see  him  laid  out  horizonally  and  surrounded  by 
flowers." 

But  there  are  rewards -a  sense  of  accomplish- 
ment and  the  joy  of  creating-  a  sense  of  helping 
others  to  participate  in  "our  glorious  science  of 
conchology"  as  Averell  put  it  so  often  in  his 
editorials  of  a  hundred  years  ago.  A  few  years 
ago  I  gathered  together  and  published  a  collec- 
tion of  interesting  articles  and  reproductions  of 
ancient  advertisements  (Abbott,  1975).  In  The 
Best  of  The  Nautilus  one  can  sense  the  spirit  of 
comradship  among  our  early  contributors.  The 
Nautilus  is  for  both  the  lieginning  scientist,  the 
enthusiastic  amateur  with  new  discoveries,  as 
well  as  for  the  accomplished  malacologist  on  the 
"cutting  edge  of  the  science"  and  immersed  in 
"the  state  of  the  art."  It  is  still  a  joint  and 
cooperative  effort.  Let's  hope  that  The  Nautilus 
goes  on  for  another  100  years! 

-R.  Tucker  Abbott,  Melbourne,  Florida 

LITERATURE  CITED 

Abbott,    R.    Tucker.    1958.    From    the    Pilsbry    Chair    of 

Malacology.  The  Nautilus  71(3):100-103. 
(editor).  1975.  The  Best  ofTheNnutilus.  280  pp., 

American  Malacologists,  Inc.,  Greenville,  Delaware. 
1979.  Bernadine  Barker  Baker -A  Tribute  upon 


her  Retirement.  The  Nnutiliis  93(2-3):ii. 
1983.   Charles   B.    Wurtz-An   Obituary.   The 


Nautilus  97(1  ):43.  portrait. 


f331    RENEWAL  OF  SUBSCRIPTION 


Enclosed  find  Two  Dollars  for  one  year's  subscription  to  THE  NAUTILUS, 
234  Berkeley  Street,  Boston,  Mass. 


Name, 


Address, 


Notice.— The  rules  now  governing  second-class  matter  do  not  permit  publications  to  be  mailed  to 
subscribers  in  arrears. 


8        THE  NAUTILUS 


January  31,  1986 


\'(il.  10(1(1) 


Abbott,    R.   Tuckor  ami   Chas.    B.    Wurtz.    1971.    Horace 

Burrington  Baker,  1889-1971.  (An  ol)ituary).  The  Nautilus 

85(l):l-4.  3  portraits. 
Avereli,  William   D.  (1899).   Pospectus  for  Tlie  Nautilus. 

1  page,  privately  printed.  Mt.  Airy,  Philadelphia,  Penna. 
Baker,    H.    Burringti)n.    1958.    Henry    Augustus    Pilsbry, 

1862-1957-An    Obituary.     The    Nautilu.'<    71(3):73-83, 

3  portraits. 
1958a.    The    Pilsbry    Nautilus.    The   Nautilu.s 

71(30):112-115. 


Gray,  Arthur  F.  1933.  Charles  WillLson  Johnson,  18()3-1932. 

(An  obituary).  The  Nautilus  46(4):  129- 136,  portrait. 
.Jutting,  W.  S.  S.  Van  B.  and  C.  0.  van  Regteren  Altena. 

1958.  A  list  of  Printed  Malacological  Journals.  Basteria 

22:10-15,  1  graph. 
Pilsbry,  H.  A.  and  C.  W.  Johnson.  1893.  To  Conchologists 

(a  piea  for  support).  The  Nautilus  7(3):25-26. 
Wurtz,    Charles    B.    1958.    Dr.    Pilsbry   and   Fresh-water 

Mollusca.  The  Nautilus  71(3):84-87. 


TOM  PULLEY  AND  THE  TRAIN 


Thomas  Edward  Pulley,  born  October  15,  1916,  Director  of  the  Houston 
Museum  of  Natural  Sciences  since  1957,  and  a  malacological  student  of  the  late 
William  J.  Clench  of  Harvard,  passed  away  on  Nove^riber  19,  1985,  at  the  age  of 
69.  In  preparation  for  the  100th  anniversary  volume  of  The  Nautilus,  which  in 
part  is  dedicated  to  Dr.  Clench,  I  asked  Tom  to  write  a  personal  anecdote  that 
typified  his  mentor.  If  my  readers  will  recall,  in  my  "Farewell  to  Bill  Clench" 
(The  Nautilus  9S(2):55-58),  Bill  drew  a  huge  arrow  in  the  sandy  beach  for  his 
students  that  pointed  to  a  waiting  train.  Tom  Pulley  has  now  caught  up  with  the 
other  students,  and  here  is  Tom's  personal  account  of  The  Train  written  in  July 
198U.  He  asked  that  this  last  picture  of  Bill  Clench  be  included- (RT A). 


Everyone  who  knew  Bill  was  aware  of  his 
capacity  for  enthusiasm  over  a  new  shell.  We 
who  worked  with  him  at  the  MCZ  often  wit- 
nessed the  thrill  of  anticipation  with  which  he 
opened  each  new  package  of  shells  and  spread 
out  the  contents  on  the  wide  brown  table.  It  was 
almost  like  watching  the  bright-eyed  wonder  of 
a  small  child  at  his  first  real  Christmas. 

But  Bill  had  the  same  happy  way  of  appreciat- 
ing most  of  the  other  simple  pleasures  of  life.  He 
and  Julia  once  invited  me  and  my  wife  and 
5-year-old  son  to  a  Sunday  afternoon  picnic;  Bill 
would  grill  the  hamburgers. 

When  we  arrived  at  his  chosen  site  I  was  a  bit 
surprised  to  see  that  we  were  on  a  nice  grassy 
spot,  but  it  was  on  the  railroad-right-of-way. 
There  were  houses  nearby,  but  they  were  mostly 
concealed  by  trees.  I  had  known  that  Bill  liked 
trains,  and  I  soon  learned  that  we  were  here  to 
wave  at  the  afternoon  train  when  it  passed  by. 
We  lighted  the  charcoal  and  opened  some  beer 
and  cokes. 

When  the  coals  were  ready.  Bill  grilled  the 
hamburgers  and  they  were  delicious.  As  we 
were  finishing  I  saw  that  Bill  key)t  glancing  at 
his  watch.  Soon  we  heard  the  faint  whistle  of 


William  J.  Clench,  age  8G,  in  California. 


Vol.  100(1) 


JanuarvSl,  1986 


THE  NAUTILUS        9 


one  of  the  last  steam  trains,  and  Bill  quickly 
jumped  to  his  feet.  He  strained  to  view  the  big 
engine  when  it  first  came  into  sight  around  a 
distant  bend,  and  long  before  it  was  near  us  he 
raised  both  hands  in  the  air  and  began  to  wave. 
As  the  train  approached  he  began  jumping  in 
the  air  and  waving  more  excitedly.  The  engineer 
saw  him  and  began  tooting  his  whistle  in  a  long 
series  of  short  bursts.  The  excitement  was 
catching,  and  by  the  time  the  engine  had 
reached  us  my  son  was  as  excited  as  Bill,  and  I 
was  amazed  to  find  that  I,  too,  was  jumping  and 
waving  like  a  5-year  old.  The  smiles  that  were 


exchanged  between  Bill  and  the  engineer  as  the 
train  roared  by  led  me  to  believe  that  this  must 
have  happened  many  times  before. 

I  understood  Bill's  joy  that  afternoon  as  the 
train  passed  by,  because  I  had  taken  part  in  it.  I 
later  realized  that  Bill  experienced  this  same 
kind  of  happy  pleasure  in  many  of  the  less 
dramatic  events  of  daily  life.  To  Bill,  the  thrill  of 
seeing  a  new  shell,  meeting  an  old  friend  or 
making  a  new  one,  finishing  another  number  of 
Johnsonia,  or  even  telling  one  of  his  old  bad 
jokes  was  comparable  to  his  pleasure  that  day  by 
the  railroad  tracks.  -  Tom  Pulley,  Julv  1984 


JOSEPH  ROSEWATER  (1928-1985) 
A  TRIBUTE  AND  BIBLIOGRAPHY 


Harald  A.  Rehder 

Smithsonian  Institution 
Washington,  D.C.  20560 


On  March  20,  1985,  malacologists  the  world 
over  lost  an  eminent  and  treasured  colleague, 
and  those  close  to  him  a  dear  friend.  His  un- 
timely death  left  a  void  in  the  ranks  of  malacolo- 
gists that  will  be  hard  to  fill,  and  his  associates 
at  the  Smithsonian  still  find  it  difficult  to  believe 
that  we  will  not  see  him  at  any  moment  leaving 
his  room  or  coming  around  the  corner  in  the 
halls  or  in  the  range  of  the  Division  of  Mollusks. 

Joe  was  born  on  September  18,  1928  in  Clare- 
mont.  New  Hampshire,  in  the  Connecticut  River 
Valley,  the  only  child  of  Joseph  and  Alice 
Tipping  Rosewater.  His  mother  had  come  to 
Claremont  from  the  Rosewater  home  in  Queens, 
Long  Island,  New  York,  to  be  with  her  parents 
at  this  time. 

Joe's  father  was  an  accountant  with  the  City 
of  New  York,  and  his  mother  had  been  a  Navy 
nurse  in  World  War  I.  When  Joe  was  eight  years 
old  his  father  died.  Joe  graduated  in  June  1946 
from  John  Adams  High  School  in  South  Ozone 
Park,  Long  Island,  two  months  after  the  death 
of  his  mother.  Joe's  mother,  before  her  death, 
had  appointed  her  brother  Charles  to  be  Joe's 
legal  guardian,  and  therefore  Joe  moved  to  his 
uncle's  home  in  Claremont,  New  Hampshire. 

In  the  fall  of  1946  Joe  entered  the  University 


of  New  Hampshire  at  Durham,  graduating  in 
1950.  As  a  boy  he  had  had  an  interest  in  animals, 
and  had  kept  an  aquarium  in  his  room.  His 
special  interest  in  mollusks  began  apparently  in 


FIG.    1.  .Jo.suph  RusuwaliT  ui  a  I'ui-fiit  iiitVirnial  puse. 


10      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


his  junior  year  at  the  University  when  his 
zoology  professors,  Emery  F.  Swan  and  George 
M.  Moore,  persuaded  him  to  undertake  a  mollus- 
can  project  for  a  master's  degree.  This  is 
evidenced  by  the  fact  that  Joe  states  that  his 
first  field  work  started  in  1949  with  collecting 
mollusks  in  New  England. 

Joe  began  working  on  his  Master's  degree  in 
the  fall  of  1950  but  his  plans  were  interrupted  by 
the  arrival  of  a  draft  notice  in  February  1951. 
That  month  he  married  Mary  Carlson  of  Gilman- 
ton,  New  Hampshire,  and  in  May  he  was  formal- 
ly drafted  and  went  to  Fort  Meade,  Maryland, 
for  basic  training.  In  the  fall  he  was  sent  to  Fort 
Sam  Houston,  Texas,  to  receive  further  training 
as  a  medical  technician  in  radiology.  In  early 
March  1952  Joe  was  sent  overseas  to  Germany 
where  until  April  1953  he  was  a  medical  techni- 
cian in  radiology  in  the  97th  General  Hospital  in 
Frankfurt-am-Main. 

After  his  return  from  Germany  and  after  leav- 
ing the  service,  Joe  worked  as  an  x-ray  tech- 
nician at  the  Sacred  Heart  Hospital  in 
Manchester,  New  Hampshire,  and  as  an  instruc- 
tor in  biology  at  Mount  St.  Mary's  College  in 
nearby  Hooksett  until  May  1955.  That  summer 
he  enrolled  at  the  University  of  New  Hamp- 
shire, attending  the  summer  school  and  then  the 
academic  year  1955-56,  receiving  a  Master's 
degree  in  June  1956. 

In  the  fall  of  1956  he  entered  Harvard  Univer- 
sity as  a  graduate  student  to  work  under  Dr. 
William  J.  Clench  in  malacology.  Here,  in  the 
company  of  Richard  I.  Johnson,  Richard  W. 
Foster,  Robert  Robertson,  Arthur  H.  Clarke 
and  Arthur  S.  Merrill,  he  spent  four  happy  and 
fruitful  years  under  the  genial  and  caring 
tutelage  of  Bill  Clench  and  Ruth  Turner. 

In  the  summer  of  1957  Joe  came  to  Washing- 
ton and  spent  three  months  in  the  Division  of 
Mollusks  of  the  U.  S.  National  Museum  as  a 
Summer  Intern,  working  on  the  family  Pleuro- 
ceridae  in  connection  with  his  doctoral  studies. 
In  the  course  of  his  studies  he  reorganized  the 
North  American  members  of  the  family 
Thiaridae  in  the  museum  collection. 

The  following  summer  Joe  accompanied  Bill 
Clench  on  a  field  trip  through  Kentucky,  Ten- 
nessee and  Georgia,  collecting  and  studying  the 
freshwater  faunas  of  the  area. 

During  his  years  at  Harvard  Joe  was  a  Teach- 


ing Fellow  in  Biology,  and  in  his  last  year  he 
held  a  position  as  Curatorial  Assistant  in  the 
Department  of  Mollusks.  In  the  fall  of  1959, 
Joe's  final  year  at  Harvard,  Bill  Clench  and  Joe 
were  beginning  to  plan  for  Joe's  immediate 
future  after  he  received  his  degree  in  the  coming 
June,  and  Bill  wrote  to  me  regarding  a  possible 
opening  at  the  National  Museum.  For  a  year  or 
more  we  had  on  file  a  job  application  form  that 
Joe  had  filled  out  for  a  position  in  the  Division, 
awaiting  the  time  when  we  would  get  approval 
to  hire  an  additional  staff  member.  At  this  time 
the  opportunity  opened  up  to  hire  a  malacologist 
under  a  grant  that  the  National  Museum  had 
received  from  the  Office  of  Naval  Research  and 
Atomic  Energy  Commission  (ONR-AEC)  to 
work  on  the  marine  faunas  of  the  Pacific, 
especially  that  of  the  Marshall  Islands- Bikini 
and  Enewetak. 

This  position  was  offered  to  Joe  and  he  started 
working  on  January  1,  1960.  In  the  year  and 
nine  months  that  he  was  on  this  contract  he 
helped  identify  and  arrange  many  lots  of  Indo- 
Pacific  mollusks,  and  spent  several  days  a  week 
reorganizing  and  expanding  the  collections  of 
Indo-Pacific  marine  mollusks,  bringing  the 
classification  up  to  date  and  adding  thousands  of 
lots  to  the  collection.  In  August  1961  Joe  made 
his  first  visit  to  the  Pacific  attending  the  Tenth 
Pacific  Science  Congress  in  Hawaii. 

On  October  2,  1961  Joe  joined  the  staff  as 
Associate  Curator,  and  for  the  next  twenty- 
three  and  a  half  years  was  a  valued  and  impor- 
tant member  of  the  scientific  staff  of  the 
National  Museum  of  Natural  History. 

With  the  addition  of  Joe,  the  staff  of  the  Divi- 
sion of  Mollusks  was  once  more  at  full  strength, 
and  this  fact  seemed  to  infuse  new  energj'  into 
the  divisional  activities,  stimulated  in  part  no 
doubt  by  Joe's  enthusiasm  in  his  new  position. 
Recorded  in  the  annual  report  for  that  year  was 
a  notable  increase  in  the  number  of  specimens 
distributed  in  exchange,  lent  for  study  and  iden- 
tified for  correspondents. 

Although  Joe's  doctoral  dissertation  dealt 
with  a  freshwater  mollusk,  marine  mollusks 
were  his  first  love,  as  he  told  me  in  a  letter  he 
wrote  when  I  offered  him  the  opportunity  of 
coming  to  the  Smithsonian  on  the  ONR-AEC 
grant.  He  had  just  finished  preparing  for  the 
cataloguer  the  last  of  the  marine  collection  of 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      11 


the  old  Boston  Society  of  Natural  History  that 
had  been  turned  over  to  the  Museum  of  Com- 
parative Zoology',  and  was  in  the  midst  of  incor- 
porating Tucker  Abbott's  Philippine  marine 
shells  into  the  collection.  When  he  came  to  work 
in  the  Division  under  the  grant,  sorting  and 
classifying  the  Indo-Pacific  moUusks  was, 
therefore,  a  familiar  task.  After  he  and  Ruth 
Turner  had  finished  the  western  Atlantic  Pin- 
nidae  they  had  planned  to  monograph  the  Indo- 
Pacific  species  of  the  family.  Now  Ruth  sug- 
gested that  Joe  do  the  paper  alone,  and  this  he 
did.  He  finished  the  project  and  it  was  published 
before  the  end  of  his  contract.  Following  this  he 
began  studying  the  family  Tridacnidae  and  the 
Indo-Pacific  Littorinidae,  both  of  which  were 
eventually  published  in  1965  and  in  1970/72.  He 
continued  his  interest  in  these  three  families  and 
published  several  papers.  He  was  working  on  a 
study  of  the  Eastern  Pacific  members  of  the 
Littorinidae  at  the  time  of  his  death,  and  also 
had  underway  a  study  of  the  bivalve  family, 
Periplomatidae. 

During  his  years  in  the  Division  of  Mollusks 
Joe  went  on  several  expeditions  and  field  trips. 
In  February  and  March  1963  he  spent  six  weeks 
at  the  Enewetak  Marine  Biological  Laboratory 
of  the  Atomic  Energy  Commission,  and  that 
winter  he  spent  three  months  in  the  western 
Indian  Ocean  on  the  research  vessel  Te  Vega  as 
part  of  the  International  Indian  Ocean  Expedi- 
tion. In  August  and  September  1966  Joe  was  in 
Australia  spending  most  of  the  time  carrying  on 
field  work  in  Western  Australia  with  Barry  C. 
Wilson,  and  in  May  and  June  1970  he  was  a 
member  of  the  Mariel  King  Memorial  Moluccas 
Expedition  on  the  R/V  Pele.  again  together  with 
Barry  Wilson.  In  April  1974  he  spent  two  weeks 
in  Tunisia  at  the  Mediterranean  Marine  Sorting 
Center,  and  in  July  1976  he  was  a  member  of  a 
party  from  the  Smithsonian  that  spent  ten  days 
making  a  survey  of  the  littoral  fauna  of  Ascen- 
sion Island.  Later  he  made  several  trips  to 
Panama,  and  to  the  Pacific  coast,  including 
Alaska,  in  connection  with  his  study  on  the  Lit- 
torinidae of  that  fauna.  His  last  trips  away  from 
Washington  were  to  the  Harbor  Branch  Labora- 
tory at  Fort  Pierce,  Florida  where  for  several 
years  he  carried  on  field  studies  on  Periploma 
and  the  Littorinidae. 

Joe  was  a  member  of  the  American  Malacolo- 


gical  Union  from  1957  on  and  was  its  President 
in  1969.  He  was  also  a  member  and  past  presi- 
dent of  the  Biological  Society  of  Washington, 
the  Society  of  Systematic  Zoology  of  which  he 
was  treasurer  1963-1966,  and  the  Paleontolo- 
gical  Research  Institute,  Ithaca,  New  York.  He 
was  a  member  of  the  National  Capital  Shell  Club 
and  its  president  in  1965.  He  served  as  a  Con- 
sulting Editor  of  The  Nautilus  from  1972  to 
1984,  and  contributed  17  articles  to  that  journal. 

In  the  Museum  he  represented  the  Depart- 
ment of  Invertebrate  Zoology  on  several  com- 
mittees-the  Professional  Advancement  Evalu- 
ation Committee,  the  Council  of  the  Senate  of 
Scientists,  and  the  Advisory  Committee  on  the 
Naturalist  Center.  On  the  Departmental  Collec- 
tions Advisory  Committee  he  represented  the 
Division  of  Mollusks. 

Joe  is  survived  by  his  wife  Mary  Carlson 
Rosewater  and  three  children,  Katherine  Louise 
Rosewater  (Waitt)  of  Sandown,  New  Hamp- 
shire, Gail  Ann  Rosewater  of  Rockville,  Mary- 
land, and  Carl  Joseph  Rosewater  of  Kensington, 
Maryland,  and  a  granddaughter,  Megan 
Rosewater- Waitt. 


Bibliography  of 
Joseph  Rosewater 

1957.  A  comparative  study  of  development  in  the  Peri- 
winkles. Abstracts  of  student  research  projects.  Woods 
Hole  Embryology  Counie.  1957:14-15,  1  table. 

1958a.  The  Family  Pinnidae  in  the  Western  Atlantic. 
Johnsonia  3(38):285-326,  23  plates  (with  Ruth  Turner). 

1958b.  Book  Review:  Olsson,  A.,  et  al.  Pliocene  Mollusks  of 
Southern  Florida,  etc.  Johnsonia  3(38):328. 

1959a.  Mollusks  of  the  Salt  River,  Kentucky.  The  Nautilus 
73(2):57-63. 

1959b.  Intertidal  stranding  of  Clione  limacina  in  Massa- 
chusetts. TheNautUux  73{2):76-n. 

1959c.  Calvin  Goodrich,  a  bibliography  and  catalogue  of  his 
species.  Occasional  Papers  on  MoUusks.  Department  of 
Mollusks.  Museum  of  Comparative  Zoology.  Hanmrd 
University.  2(24):  189-208,  1  photograph,  1  plate. 

1959d.  A  Pleurocerid  index  to  past  collecting  in  the  Ohio 
and  Tennessee  River  Basins  with  some  thoughts  for  the 
future.  Annual  Report  American  Malacological  Union 
for  1958,  Bull.  25:23-24. 

1960a.  Egg  mass  and  gross  embryology'  of  Pleurocera 
canalicutata  (Say,  1821).  Annual  Report  .\merican  Mala- 
cological Union  for  19.59,  Bull.  26:10-12. 

1960b.  Proposed  further  use  of  the  Plenary  Powers  in  the 
case  of  the  Generic  name  Pleurocera  Rafinesque,  1818 
(Class  Gastropoda).  Z.M.(S)83.  Bulletin  of  Zoological 
NoTnenclature  17(6-8):171-172. 

1961a.  Preliminary  observations  on  the  nervous  systems  of 


12      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


some  Pleuroceridae.  Annunt  Report  American  Mahi- 
cological  Union  for  1961,  Bull.  28:10-11. 

1961b.  Comments  on  the  proposal  to  place  the  Generic  name 
Gari  Schumacher,  1817,  on  the  Official  List,  Unemended, 
Z.N.(S)1461.  Bulletin  of  Zoological  Nomenclature 
18(5):303. 

1961c.  The  Family  Pinnidae  in  the  Indo-Pacific.  Indo- 
Pacific  Mollwica  l(4):17.'5-226,  pis.  135-171,  figs.,  maps. 

1963a.  Resistance  to  dessication  in  dormancy  by  Tectariux 
muricatuit.  The  Nautilus  76(2):111. 

1963b.  An  effective  anesthetic  for  giant  clams  and  other 
mollusks.  Turtux  News  (Chicago)  41(12):300-302,  2  figs. 

1963c.  Problems  of  species  analogues  in  world  Littorinidae. 
Annual  Report  Anifrican  Malacolociical  Union  for  1963, 
Bull.  30:.5-6. 

196.5a.  The  Family  Tridacnidae  in  the  Indo  Pacific.  Indo- 
Pncific  Mollusra  l(6):347-396,  pis.  263-293. 

19651).  Studying  living  Tridacnidae  in  the  Marshall  Islands. 
Annual  Report  American  Malacological  Union  for  1965, 
Bull.  32:18-19. 

1966a.  Reinstatement  of  Melarhaphe  Menke,  1828.  The 
Nautilus  80(2):37-38. 

1966b.  The  Giant  Clams.  Australian  Natural  History  1.5(8): 
250-256.  5  figs. 

1967a.  Indo- West  Pacific  Littorinidae.  Aurnial  Report 
American  Malacological  Union  for  1966,  Bull.  33:27. 

1967b.  Book  Review:  Halstead,  B.  W.  1965.  Poisonous  and 
Venomous  Marine  Animals  of  the  World,  vol.  1,  Inverte- 
brates. Science  156(3776):795  (with  M.  L.  .Jones,  R.  B. 
Manning,  D.  L.  Pawson,  and  K.  Ruetzler). 

1968a.  The  Zoological  Taxa  of  William  Healey  Dall.  Bulletin 
U.  S.  National  Museum  287:1-427  (with  K.  J.  Buss  and 
F.  A.  Ruhoff). 

1968b.  Itinerary  of  the  Voyage  of  H.  M.  S.  Blossom.  The 
Veliger  10(4):'350-352. 

1968c.  Book  review:  Hyman,  L.  H.  1967.  The  Invertebrates, 
vol.  VI.  Mollusca  1.  Science  160(3823):62-63. 

1968d.  Book  review:  An  English-Classical  Dictionary  for  the 
use  of  Taxonomists.  Systematic  Zoology  17(3):334. 

1968e.  Notes  on  Periplomatidae  (Pelecypoda:  Anomalodes- 
mata)  with  a  geographical  checklist.  Annual  Report 
American  Malacological  Union  for  1968,  Bull.  35:37-39. 

1969a.  (iross  anatomy  and  classification  of  the  commensal 
gastropod,  CaledonieUa  montrouzieri  Souverbie,  1869. 
The  Veliger  ll(4):345-350,  text  figs.  1-4,  pi.  55. 

1969b.  George  Mitchell  Moore.  1906-1968.  [Obituary].  The 
Nautilus  9,ZC[):\\\. 

1969c.  Comment  on  a  review  by  ,J.  D.  Rising  of  R.  E.  Black- 
welder,  Taxonomy,  a  Text  and  Reference  Book.  Systema- 
tic Zoology  18(3):361-362. 

1969d.  Malacological  collections-development  and  manage- 
ment. In:  Natural  History  Collections,  Past- Present - 
Future,  Symposium  of  the  Biological  Society  of  Washing- 
ton. Proceedings  of  the  Biological  Society  of  Washington 
82:663-670. 

1970a.  The  Family  Littorinidae  in  the  Indo-Pacific.  Part  I. 
Subfamily  Littorininae.  Indo-Pacific  Mollusca  2(11): 
417-506,  64  plates. 

1970b.  Monoplacophora  in  the  South  Atlantic  Ocean.  Science 
167(3924):1485-1486,  fig. 

1970c.  Another  record  of  insect  dispersal  of  an  Ancylid 
snail.  TheNautihis  83(4):144-145. 


I97()d.  The  role  of  the  professional  in  Malacologj-.  Pitts- 
hurgh  Shell  Club  Bulletin  5:12-13. 

1970e.  Book  Review  of:  .Johnstone,  Kathleen  Yerger.  Col- 
lecting Seashells.  Grosset  and  Dunlap.  Illus.,  198  pp. 
Atlantic  Naturalist  25(4):187-188. 

1970f.  Discussion  of:  R.  T.  Abbott,  Eastern  Marine  Mol- 
lusks. In:  American  Malacological  Union  Symposium  on 
Rare  and  Endangered  Mollusks.  Malacologia  10(1):49. 

1972a.  The  Family  Littorinidae  in  the  Indo-Pacific.  Part  II. 
The  Subfamilies  Tectariinae  and  Echinininae.  Indo-Pacific 
Mollusca  2(12):507-534,  21  plates. 

1972b.  Book  Review:  Wilson,  Barry  R.  and  Keith  Gillett. 
1971.  Australian  Shells.  National  Capital  Shell  Club 
Newsletter  February  1972,  2  pp. 

1972c.  Book  Review:  Brost,  F.  B.  and  R.  D.  Coale.  1971. 
A  guide  to  Shell  Collecting  in  the  Kwajalein  .-Xtoll. 
National  Capital  Shell  Club  Newsletter  February  1972, 
2  pp. 

1972d.  The  Amphi-Atlantic  distribution  of  Liltorina  melea- 
gris.  The  Nautilus  86(2-4):67-69,  plate,  map.  table  (with 
G.  .J.  Vermeij). 

1972e.  Teratological  Littorina  scahra  angulifera.  The 
Nautilus  86(2-4):70-71,  plate. 

1972f.  Life  spans  of  Mollusca.  In:  Altman,  P.  L.  and  D.  S. 
Dittmer.  1972.  Biology  Data  Book,  edition  2,  vol.  1, 
pp.  234-235.  Federation  of  American  Societies  for  Experi- 
mental Biologj',  Bethesda,  Maryland. 

1973a.  Book  Review:  Wilson,  B.  R.  and  Keith  Gillett.  1971. 
Australian  Shells.  The  Quarterly  Rerieiv  of  Biology 
48(l):42-43. 

1973b.  Book  Review:  Graham,  Alastair.  1971.  British  Proso- 
branch  and  other  Operculate  Gastropod  Molluscs.  The 
Quarterly  Review  of  Biology  48(1):43. 

1973c.  A  Source  of  Authors  and  Dates  for  Family  Names  of 
Gastropods.  The  Veliger  16(2):243. 

1973d.  Replacement  pages  for:  The  Family  Littorinidae  in 
the  Indo-Pacific,  Part  II,  the  Subfamilies  Tectariinae  and 
Echinininae  (Indo-Pacific  Mollusca  2(12):509-516). 
Indo-Pacific  Mollusca  3(14):63-70. 

1973e.  More  on  Penis  Shedding  Among  L('/ion)ja.  Neir  York- 
Shell  Club  Notes,  no.  196,  November  1973,  p.  7. 

1974a.  Studies  on  Ascension  Island  Marine  Mollusks.  Bulk- 
tin  American  Malacological  Union  for  1973,  pp.  30-32. 

1974b.  Phylogeny  of  Littorinidae.  Littorinid  Tidings. 
Occasional  Newsletter  of  Littorinidae  Research  Grotip, 
no.  1,  pp.  10-11. 

1975a.  An  Annotated  List  of  the  Marine  Mollusks  of  Ascen- 
sion Island,  South  Atlantic  Ocean.  Smithsonian  Contri- 
butions to  Zoology  189:1-41,  figs.  1-24,  3  tables. 

1975b.  The  Marine  Commensal  Gastropod,  CaledonieUa 
mxmtrouzieri  (Prosobranchia:  Hipponicacea)  in  Thailand. 
The  Nautilus  $%iyM. 

1975c.  Mollusks  of  Gatun  Locks,  Panama  Canal.  Bulletin 
American  Malacological  Union  for  1974,  pp.  42-43. 

1975d.  Book  Review:  R.  Scase  and  E.  Storey.  1975.  The 
World  of  Shells.  National  Capital  Shell  Chih  Neu'sletter. 
September,  1975.  p.  10. 

1976a.  William  Healey  Dall,  The  Legacy  He  Left  for  Mala- 
cology. Bulletin  American  Malacological  Union  for  1975, 
pp.  4-6. 

1976b.  The  NMNH-STRI  Survey  of  Panama  1971-1975. 
Bulletin    American    Malacological    Union    for    1975, 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      13 


pp.  48-50. 
1976c.  Letter  U>  the  Editor.  AiistnilniH  Slifll  ,\eirx  No.  Hi, 

p.  2. 
1976(1.   Book  Review:  Ruth  Fair.  Shell  Collectors'  Cuide. 

National  Capital  Shtil  Cliili  Ni-iivlettrr.  Septemlier  1976, 

pp.  11-12. 
1976e.  Application  to  use  Plenary  Power  to  designate  the 

type  of  Pleurocera.  Bulletin  of  Zmilogiral  Nnnirnrlatiirr 

33(2):10.->-n3. 
1977.  Bicentennial  Booty.  National  Capital  Shrll  CluliNcivs- 

letter.  February  1971,  p.  8. 
1978a.  Zoogeography  of  West  African  Littorinidae.  Bulletin 

Ameriran  Malacological  Union  for  1977.  pp.  .31-34. 
1978b.  A  Case  of  Double  Primary  Homonymy  in  East  Pacific 

Littorinidae.  The  Nautilus  92(3):123-12.5. 
1978c.  A  Malacological  Ecpedition  to  the  Moluccas,  National 

Geiyraphir  Society  Research  Reports  nf  1969  Projects. 

pp.  .515-523.  (With  Barry  R.  Wilson). 
1978d.  Book  Review;  Kirtisinghe.  P.   1978.  Sea  Shells  of 

Sri    Lanka.    National    Capital    Shell    Club    Newsletter. 

September  1978,  p.  11. 
1979a.  A  Reconnaissance  of  West  American  Littorinidae. 

Bulletin  American  Malacological  Union  for  1978,  p,  55. 
1977b.  Rectifications  in  the  Nomenclature  of  some  Indo- 

Pacific   Littorinidae.   Proceedings  Biological  Society   of 

Wa.-ihington  92(4):773-782.  (with  W.  F.  Ponder). 
1979c.  Further  Comments  on  the  Proposed  Designation  of  a 

Type  Species  for  Pleurocera  Rafinesque,  1818.  Z.N.(S)83. 

Bulletin  Zoological  Nommclature  36(3):139-146.   (with 

J.  P.  E.  Morrison,  A.  H.  Clarke,  G.  M.  Davis,  and  C.  B. 

Stein). 
1980a.   Predator  Boreholes  in  Periploma   margaritaceuni 

with  a  brief  survey  of  other  Periplomatidae  (Bivalvia: 

Anomalodesmata).  The  VV/7>r  22(3):248-251. 
1980b.  A  Close  look  at  Littorina  Radulae.  Bulletin  Ameri- 
can Malacological  Union  for  1979,  pp.  5-8,  8  figs. 
1980c.  Subspecies  of  the  Gastropod  Littorina  .•icahra.  The 

Nautihis  94(4):158-162,  12  figs, 
1980d,  Book  Review:  Melvin,  A.  G.  and  Melvin,  L.  S.  1980. 

1000  World  Sea  Shells,  Rare  to  Common  with  Values. 

National  Capital  Shell  Club  Newsletter.  September,  p.  4. 
1981a.  The  Family  Littorinidae  in  Tropical  West  Africa. 


Atlantule  Reports  13:7-48,  6  plates,  2  tables. 
1981b.  A  New  Aid  to  Taxonomic  Research  on  Mollusks. 

Bulletin  American  Malacological  Union  for  1980,  p,  67. 
1981c.  Changes  in  Shell  Morphology  of  Post  Larval  Tri- 

dacna    gigas    Linne    (Bivalvia:    Heterodonta).    Bulletin 

American  Malacological  Union  for  1980,  pp.  45-48,  figs. 
1981(1.  Malacological  .'lourney  into  Cuba,  1980,  The  Nautilus 

95(4):1.59-162, 
1981e.  Rectifications  in  the  Nomenclature  of  some  Indo- 

Pacific  Littorinidae -II.   Proceedings  Biological  Society 

of  Washington  94(4):1233-1236. 
1982a.  A  new  species  of  Hippopus  (Bivalvia:  Tridacnidae). 

The  Nautilus  96(l):3-6,  4  figs. 
1982b.  A  new  species  of  the  Genus  Echininus  (Mollusca: 

Littorinidae:  Echinininae)  with  a  review  of  the  Subfamily. 

Proceedings    Biological    Society    of    Washington    95(1): 

67-80,  6  figs,  table. 
1982c.  Book  Review:  R.  K.  Dell,  Sea  Shore  Life.  Natiaual 

Capital  Shell  Newsletter.  No.  70,  Autumn,  p.  15. 
1983a.  Review  of  Hawaiian  Pinnidae  and  Revalidation  of 

Pinna  exquisita  Dall,  Bartsch,  and  Rehder,  1938  (Bivalvia: 

Mytiloida).  Pacific  Science  36(4):453-45S. 
1983b.  Review  of:  .lames  Graham  Cooper- Pioneer  Western 

Naturalist,  by  Eugene  Coan.  The  Nautilus  97(l):44-45. 
1983e.  Another  Bivalve-/l/)/(?-o(^j;(7  Association  with  Com- 
ments on  Adaptive  Significance  of  Oddly  Shaped  Lepto- 

nacea.  American  Malacological  BuUetiii  1:90-91. 
1984a.   A  bibliography  and  list  of  the  taxa  of  Mollusca 

introduced  by  .Joseph  P.  E.  Morrison  (December  17,  1906- 

December  2,  1983).  The  Nautilus  98(l):l-9. 
1984b.  A  new  species  of  Leptonacean  Bivalve  from  off 

Northwestern  Peru  (Heterodonta:  Veneroida:  Lasaeidae). 

The  Veliger  27(l):81-89.  6  figs.  1  table. 
1984c.  Burrowing  activities  of  Periploma  margaritaceum 

(Lamarck,  1801)  (Bivalvia:  Anomalodesmata:  Periploma- 
tidae). American  Malacological  Bulletin  2:35-40,  1  plate, 

1  table. 
1984d.  Bermuda  marine  Mollusk  type  specimens  transferred 

to  the  Smithsonian.  The  Nautilus  98(4):151-153. 
1985.  Epizoan  Communities  on  Marine  Turtles  1.  Bivalve 

and  Gastropod  Mollusks.  Marine  Ecology  6(2):127-140, 

4  figs.,  1  table  (with  .1.  Frazier  et  al.). 


14      THE  NAUTILUS 


lanuary  31,  1986 


Vol.  100(1) 


DESCRIPTION  OF  THE  HABITAT  OF  THE  ENDANGERED  MUSSEL 

PLETHOBASUS  COOPERIANUS 


Andrew  C.  Miller 
Barry  S.  Payne 

U.S.  Army  Engineer  Waterways 
Experiment  Station 

P.O.  Box  631 
Vicksburg,  MS  39180 


and  Terry  Siemsen 

Navigation  Planning  Support  Center 

P.O.  Box  59 

Louisville,  KY  40201 


ABSTRACT 

A  total  of  26  species  ofunionids,  in  addition  to  the  Asian  Clam.  Corbicula,  were 
collected  at  a  mussel  bed  on  the  Ohio  River  near  Ohnsted,  Illinois.  The  bed  was 
dominated  by  Corbicula  (1475/m^,  128.2  g/m^  of  Tissue  Dry  Mass,  TDM)  and 
Fusconaia  ebena  (66.0/m^  and  39.7lm^  of  TDM)  although  eight  other  unionids. 
Amblema  plicata,  Elliptio  dilatatus,  Leptodea  fragilis,  Ligumia  recta,  Mega- 
lonaias  gigantea,  Pleurobema  cordatum,  Potamilus  alatus,  and  Truncilla  donaci- 
formis  were  also  judged  comynon.  In  addition,  three  live  specimens  of  the  Fed- 
erally endangered  Orange-footed  Pimpleback  Mussel,  Plethobasus  cooperianus, 
were  identified.  All  appeared  healthy  and  were  adult-sized,  ranging  in  total  shell 
length  from  68  to  7U  mm.  While  there  are  no  records  of  recent  recruitment  for  P. 
cooperianus,  it  appears  that  this  mussel  bed,  which  contains  a  diverse  cammunity 
of  bivalves  in  addition  to  a  dense  population  o/' Corbicula,  provides  good  habitat 
for  this  particular  species. 


Three  live  specimens  of  the  Endangered 
Plethobasus  cooperianus  (Lea  1834),  the  Orange- 
footed  Pimpleback  Mussel,  were  found  in 
September  1983  during  a  survey  of  a  gravel  bar 
in  the  Ohio  River  near  Olmsted,  Illinois.  In  the 
summer  of  1982,  Williams  and  Schuster  (1982) 
collected  mussels  at  this  site  and  found  a  single 
live  P.  cooperianus.  However,  other  workers 
brailed  this  gravel  bar  and  found  no  live  endan- 
gered species  (Neff  &  Pearson  1980,  Williams 
1969).  The  purpose  of  this  paper  is  to  document 
the  existence  of  this  species,  and  to  provide  in- 
formation on  habitat  characteristics  and  com- 
munity structure  where  it  was  collected. 

The  historical  range  of  P.  cooperianus  in- 
cluded the  Ohio  River  from  western  Pennsyl- 
vania to  southern  Indiana;  the  Wabash  River 
below  Mt.  Carmel,  Illinois;  the  Cumberland 
River  from  Cumberland  County,  Kentucky,  to 
the  vicinity  of  Nashville,  Tennessee;  the  lower 
Clincli  River  in  Anderson  County,  Tennessee; 
and  the  Tennessee  River  from  near  Knoxville, 
Tennessee,  to  Kentucky  Lake,  Benton  County, 
Tennessee.  It  has  also  been  recorded  from  the 
Caney  Fork,   Holston,   and  French  Rivers  in 


Tennessee  and  from  the  Green  and  Rough 
Rivers  in  Kentucky.  At  present,  P.  cooperianus 
is  restricted  to  the  Ohio  and  lower  Wabash 
Rivers  where  it  is  uncommon  and  to  the  lower 
Tennessee  River  in  Alabama  and  western 
Tennessee  where  it  is  also  uncommon. 

The  shell  of  P.  cooperianus  is  up  to  87  mm 
long,  75  mm  high,  and  45  mm  wide;  it  is  heavy, 
subcircular,  and  tuberculate.  The  periostracum 
is  chestnut  colored  and  the  nacre  is  either  white 
or  faintly  pink  and  iridescent  posteriorly. 

Superficially  this  species  resembles  Q. 
pustulosa,  but  the  latter  species  is  usually 
smaller  and  has  fewer  and  relatively  larger 
tubercles  which  tend  to  be  laterally  rather  than 
radially  extended.  In  addition,  Q.  pustulosa 
exhibits  white  nacre  and  young  specimens  have 
prominent  green  rays,  whereas  in  P.  cooperia- 
nus the  rays  are  obscure  and  narrow.  The  most 
definitive  characteristic  in  P.  cooperianus  is  the 
bright  orange  viscera,  which  can  be  seen  in  live 
specimens  by  gently  prying  the  valves  apart 
(Clarke  and  Fuller  1983). 

Wilson  and  Clark  (1914)  reported  collecting 
two  gravid  females  in  June  from  the  Cumber- 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      15 


land  River;  evidently  it  is  a  summer  breeder 
although  the  host  fish  is  unknown.  Although 
ecological  data  are  virtually  non-existent,  this  is 
a  large  river  species  which  has  been  collected 
from  sand  and  gravel  substrate  in  shoals  and 
riffles. 

Methods  and  Study  Area 

On  26-29  September  1983  molluscs  were  col- 
lected from  a  reach  of  the  Ohio  River  below 
Lock  and  Dam  53  near  Olmsted,  Illinois.  The 
study  area  included  the  upper  portion  of  a 
mussel  bed  delimited  by  Williams  (1969)  and 
Williams  and  Schuster  (1982).  During  the  four- 
day  study  bivalves  were  collected  by  hand  along 
the  shore,  with  the  use  of  a  5-ft  brail  bar,  and  by 
an  experienced  shell  diver  using  underwater 
breathing  apparatus.  As  part  of  this  work  the 
diver  completed  three  30-min  qualitative 
searches,  and  collected  six  0.25-  m^  quadrat 
samples  from  each  of  four  sites  on  the  mussel 
bed.  This  project  was  undertaken  for  the  U.  S. 
Army  Engineer  District,  Louisville,  to  provide 
information  for  a  Feasibility  Report  and  Envi- 
ronmental Impact  Statement  for  the  Lower 
Ohio  River  Navigation  Project. 

The  gravel  bar  where  mussels  were  collected 
was  about  3  miles  long  and  followed  the  Illinois 
shoreline.  Substrate  consisted  of  densely  packed 
coarse  sand  and  gravel.  Particle  sizes  varied 
from  less  than  1.0  cm  to  more  than  10.0  cm  with 
the  greatest  fraction  of  total  weight  in  the  1-  to 
3-  cm  range.  Water  depths  where  P.  cooperia- 
nus  were  taken  ranged  from  approximately  3  to 
6  m  deep.  The  study  site  was  Ohio  River  Mile 
966.6  to  967.2,  approximately  4  miles  downriver 
of  Lock  and  Dam  53;  navigation  and  barge  fleet- 
ing were  taking  place  in  the  immediate  area.  No 
recent  evidence  of  sedimentation,  physical  abra- 
sion, or  propeller  wash  from  navigation  activi- 
ties, were  noted  at  this  portion  of  the  gravel  bar. 

Results 

Using  qualitative  and  quantitative  techniques, 
26  species  of  unionids,  in  addition  to  the  Asian 
Clam,  Corhicula,  were  collected  at  or  adjacent 
to  the  gravel  bar  (Table  1).  Although  nine 
species  were  judged  common,  the  most  abun- 
dant bivalves  were  Corhicula,  Amblema  plicata, 
Fusconaia  ebena,  Lampsilis  teres.  Megalonaias 
gigantea,  and  Potomilus  alatios. 


TABLE  1.  Bivalves  ccillected  at  a  gravel  bar  near  Olmsted, 
Illinois.  26-29  September  1983.  All  taxa  were  collected  alive. 

Scientific  Name 

Family  Unionidae 

Actlnonaias  carinata 

Amblema  plicata 

Cyclonaias  tuberculata 

Ellipsaria  llneolata 

Elliptio  crassidens 

Elliptio  diiatatus 

Fusconaia  ebena 

Lampsilis  ovata 

Lampsilis  teres 

Lasmigona  complanata 

Leptodea  f ragilis 

Ligumia  recta 

Megalonaias  gigantea 

Obliquaria  ref lexa 

Qbovaria  olivaria 

Flethobasus  cooperianus 

Plethobasus  cyphyus 

Pleurobema  cordatum 

Potamilis  alatus 


Quadrula  metanevra 
Quadrula  nodulata 
Quadrula  pustulosa 
Quadrula  quadrula 
TritoRonia  verrucosa 
Truncilla  donacif ormis 
Truncilla  truncata 
Family  Corbiculidae 
Corbicula  manilensis 


Presence* 

UC 
C 

nc 

FC 
FC 
UC 

c 

UC 

c 

UC 

c 
c 
c 

FC 
UC 

UC 
UC 

c 
c 

FC 
FC 
FC 
FC 
UC 
C 
DC 


*  Presence,  for  this  study: 
UC  =  Uncommon,  <  6  collected 
FC  =  Fairly  common,  6-20  collected 
C  =  Common,  >    20  collected 


A  total  of  14  species  of  unionids  were  collected 
in  24  0.25-m^  quadrat  samples  (Table  2).  Total 
unionid  density  ranged  from  28  to  124/m^  (X  = 
60).  Numerically,  F.  ebena  (X  =  46/m^)  and  Trun- 
cilla donaciformis  (X=12/m^)  dominated  the 
unionids.  Two  species,  Q.  metanevra  and  Q. 
nodulata,  judged  fairly  common  in  the  study 
area,  were  found  in  only  1  of  the  24  quantitative 
samples.  The  gravel  bar  was  numerically  domi- 
nated by  Corbicula,  with  numbers  ranging  from 
556  to  2628/m'  (X=1475).  Average  tissue  dry 
mass  (TDM)  for  Corbicula  (128.2  g/m')  was  ap- 
proximately three  times  that  for  total  unionids 
(X  =  39.7).  Because  of  their  large  size  and  high 


16      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


TABLE  2.  Bivalves  collected  in  24  0.25-ni'  quadrat  samples  at  a  gravel  bar  near  Olmsted, 
Illinois,  28-29  September  1983. 


Species 

Occurrence* 
24 

No/m^ 
1475 

TDM/m^ 

Corbiaula  manilensis 

128.2 

Total  unlonlds 

Ih 

66.0 

39.7 

Fusconaia  ebena 

24 

45.5 

27.4 

Truncilla  donaaiformis 

24 

11.5 

0.4 

Leptodea  fvagilis 

11 

2.5 

1.3 

Obliquaria  reflexa 

9 

1.8 

0.8 

Quadrula  pustulosa 

8 

1.5 

1.4 

Amblema  plioata 

6 

1.0 

3.1 

Ellipsaria   lineolata 

7 

1.3 

1.5 

Tritogonia  verrucosa 

3 

0.5 

0.4 

Lcmpsilis  ventrioosa 

6 

1.2 

0.03 

Quadrula  quadrula 

5 

0.8 

1.3 

Quadrula  metanevra 

1 

0.2 

0.1 

Quadrula  nodulata 

1 

0.2 

0.1 

Megalonaias  gigantea 

2 

0.3 

1.3 

Aatinonais  aarinata 

1 

0.2 

0.3 

*     Occurrence   -  Number   of   quadrats  with  live  organisms    (total  quadrats 
collected   =    24) . 


numbers,  F.  ebena  represented  the  majority, 
about  70%  (27.4  g/m^  of  the  total  unionid 
biomass.  Although  numerically  dominant,  T. 
donaciformis.  because  of_  its  small  size, 
represented  only  about  1%  (X  =  0.4  g/m^)  of  the 
unionid  biomass.  Based  upon  these  quantitative 
collections,  evidence  of  recent  recruitment 
(presence  of  clams  less  than  12  months  old  judg- 
ing from  their  small  size  and  unweathered  ap- 
pearance) was  observed  for  12  of  the  14  unionids 
collected  in  the  quadrats. 

The  three  specimens  of  P.  cooperianns  were 
obtained  near  Ohio  River  Mile  967.2  by  the  diver 
during  two  of  the  three  30-min  qualitative 
searches  of  the  bar.  The  diver  had  been  in- 
structed to  concentrate  on  pustulate  organisms 
or  species  judged  uncommon  by  our  earlier 
work.  On  the  second  search,  he  obtained  eight 
species,  including  one  P.  cooperianiui  and  two  Q. 
ptistulosa.  During  the  final  search  he  retrieved 
eleven  species,  including  two  P.  cooperianus  and 
two  Q.  pustulosa.  The  Orange-footed  Pimple- 
back  was  not  found  with  the  brail,  on  any  of  the 


shoreline  searches,  or  in  the  quadrat  samples. 

Conchological  data  on  the  three  live  P. 
cooperianus  are  as  follows: 

Length  (mm)             Height  (mm.)  Width  (mm) 

74                             63  37 

68                             60  38 

71                             64  42 

By  gently  prying  the  valves  apart,  it  was  ob- 
served that  soft  tissues  were  healthy  looking 
and  certainly  not  moribund. 

Discussion 

This  gravel  bar  was  dominated  both  in  num- 
bers and  biomass  by  Corbicula;  evidently  this 
had  no  effect  on  the  unionids  or  P.  cooperianus. 
In  addition,  it  appears  that  requirements  for 
food,  water  quality,  and  substrate  for  adult  P. 
cooperianus  must  differ  very  little  from  the  25 
other  unionids  inhabiting  this  bed.  In  a  study  of 
21  species  in  Michigan  streams,  Strayer  (1983) 
concluded  that  microhabitat  differences  were 
not  discernible  for  21  of  the  unionids.  In  addi- 
tion, two  other  species,  the  fairly  uncommon 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      17 


Cumberlandia  monodonta  and  the  endangered 
Lampsiliti  higginsi,  have  also  been  collected 
from  very  diverse  and  densely  populated  mussel 
beds  in  the  Upper  Mississippi  River.  Cumber- 
landia monodonta  was  usually  taken  in  samples 
where  50%  of  the  sites  had  at  least  20  species  of 
common  or  wide-ranging  mussels.  Lampsilis 
higginsi  was  in  association  with  17  or  more  com- 
mon unionids  at  50%  of  the  reported  sites  in  the 
upper  Mississippi  River  (Nelson  and  Freitag 
1979). 

Wilson  and  Clark  (1914)  judged  P.  cooperia- 
nus  to  be  "not  rare"  during  their  survey  of  the 
Cumberland  River.  However,  Neel  and  Allen 
(1964)  were  unable  to  locate  this  species  during 
a  follow-up  study  on  the  same  river.  It  is  ap- 
parent that  the  historical  range  of  P.  cooperia- 
nus  has  diminished  and  the  species  is  in  danger 
of  becoming  e.xtinct.  Some  of  the  reasons  often 
cited  for  the  loss  of  mussels  in  large  rivers  in- 
clude: sedimentation,  navigation  activities, 
pollution,  reservoir  construction  (many  with 
deoxygenated,  low  pH,  and  cold  water  releases), 
and  loss  of  fish  hosts  (Fuller  1974). 

Conversion  of  large,  free-flowing  rivers  such 
as  the  Ohio  to  navigable  waterways  altered  the 
habitats  of  many  benthic  organisms.  However, 
some  extensive  areas  with  suitable  substrate, 
flow,  water  quality,  and  nutrients  still  exist 
which  support  recruiting  unionid  communities. 
For  the  endangered  P.  cooperianus,  the  studied 
gravel  bar  on  the  Ohio  River  appears  to  meet 
requirements  necessary  for  maintenance  of 
adults. 

The  range  reduction  for  P.  cooperianus  is 
probably  related  to  species-specific  problems  of 
reproduction  or  recruitment.  Conversion  of  the 
Ohio  River  from  a  freely-flowing  river  to  a  con- 
trolled waterway  probably  modified  the  avail- 
able habitats  and  could  have  had  detrimental 
effects  on  host  fish.  However,  it  is  also  possible 
that  this  species  is  slightly  more  sensitive  to 
altered  habitats  than  the  other  thick-shelled 
unionids  at  the  mussel  bed.  Perhaps  conditions 
have  degraded  to  a  point  that  P.  cooperianus 
can  maintain  itself  but  does  not  possess  energy 
reserves  necessary  for  production  of  adequate 
numbers  of  glochidia. 

Similar  problems  may  also  be  blamed  for  loss 
of  species  in  the  genus  Dysnamia,  which  were 
once  commonly  collected  in  riffles  or  shoals  in 


large  rivers  (Stansbery  1970).  Riffle  and  shoal 
habitats  are  now  almost  non-existent  in  large 
rivers  in  North  America;  their  disappearance 
could  be  blamed  for  loss  of  recruiting  popula- 
tions oi  Dysnotnia  and  Plethobasus. 

A  possible  solution  for  long-term  maintenance 
of  P.  cooperianus  is  artificial  propagation  (Isom 
and  Hudson  1982)  and  translocation  of  im- 
mature forms  to  suitable  habitats.  Projects  such 
as  these  will  be  important  for  separating  direct 
effects  of  habitat  alteration  from  indirect  effects 
such  as  reduction  in  the  availability  of  the  host 
fish. 

Acknowledgments 

Funds  for  this  work  were  provided  by  the 
Navigation  Planning  Support  Center,  U.  S. 
Army  Engineer  District,  Louisville  in  Louisville, 
Kentucky. 

LITERATURE  CITED 

Clarke,  A.  H.,  and  Fuller,  S.  L.  H.  1983.  A  Field  Guide  to  the 
Endangered  Mussels.  Technical  Report  in  Preparation, 
U.S.  Army  Engineer  Waterways  Experiment  Station, 
Vicksburg,  Miss. 

Fuller,  S.  L.  H.  1974.  Clams  and  Mussels  (Mollusca: 
Bivalvia),  In:  Hart,  C.  W.,  Jr.,  and  S.  L.  H.  Fuller,  eds.. 
Pollution  Ecology  of  Fresh-Water  Invertebrates.  Academic 
Press,  New  York. 

Isom,  B.  G,,  and  Hudson,  R.  G.  1982.  In  vitro  Culture  of 
Parasitic  Freshwater  Mussel  Glochidia.  The  Nautilus 
96(4):147-151. 

Neel,  J.  K.,  and  Allen,  W.  R.  1964.  The  Mussel  Fauna  of  the 
Upper  Cumberland  Basin  Before  Its  Impoundment.  Mala- 
rologia  1:427-459. 

Neff,  S.  E.,  and  Pearson,  W.  1980.  Aquatic  Ecology  and 
Water  Quality  Studies  for  the  Lower  Ohio  River  Naviga- 
tion Study,  Unpublished  report  submitted  to  the  Louisville 
District  Office,  Louisville,  Ky. 

Nelson,  D.  A.,  and  Freitag,  T.  M.  1979.  Ecology,  Identifica- 
tion, and  Recent  Discoveries  of  Higgin's  Eye  {Lampsilis 
higginsi].  Spectacle  Case  (Cumberlandia  monodonta).  and 
Fat  Pocketbook  (Potamiius  capax)  Mussels  in  the  Upper 
Mississippi  River.  In:  Rasmussen,  J.  L.,  ed.,  Proceedings  of 
the  UMRCC  Symposium  on  Upper  Mississippi  River 
Bivalve  Mollusks.  J.  L.  Rasmussen,  ed.  pp.  120-145. 

Stansbery,  D.  H.  1970.  Eastern  Freshwater  Mollusks  (I)  The 
Mississippi  and  St.  Lawrence  River  Systems.  Malacologia 
10(l):9-22. 

Strayer,  D.  L.  1983.  Notes  on  the  Microhabitats  of  Unionid 
Mussels  in  Some  Michigan  Streams.  Amer.  Midi  Nat. 
106(2):411-415. 

Wilson,  C.  B.,  and  Clark,  H.  W.  1914.  The  Mussels  of  the 
Cumberland  River  and  Its  Tributaries.  Report  U.  S.  Com- 
mission Fisheries  for  1912  and  Spec.  Papers,  pp.  1-63 
(separately  issued  as  Bureau  Fisheries  Document  No.  781). 

Williams,  J.  C.   1969.  Mussel  Fishery  Investigation,  Ten- 


18      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


nessee,  Ohio,  ;ind  (Sreen  Rivers.  Final  Report.  State  of 
Kentucky  Project  Number  4-19-R,  Report  to  U.  S.  Fish 
and  Wildlife  Service,  107  pp. 
Williams,  J.  C,  and  Schuster,  G.  A.  1982.  Fresh-Water 
Mussel  Investigations  of  the  Ohio  River,  Mile  317.0  to  Mile 


981.0,  Report  Submitted  to  the  U.  S.  Army  Engineer  Dis- 
trict, Louisville;  Kentucky  Department  of  Fish  and  Wild- 
life Resources;  Division  of  Fisheries,  U.  S.  Fish  and  Wild- 
life Service,  Areas  3  and  4. 


CHARACTERIZATION  OF  A  BIVALVE  COMMUNITY  IN  THE 
TANGIPAHOA  RIVER,  MISSISSIPPI 


Andrew  C.  Miller 
Barry  S.  Payne 

Environmental  Laboratory 

U.S.  Army  Engineer  Waterways  Exp.  Station 

P.O.  Box  631,  Vicksburg,  MS  39180 


and  David  W.  Aldridge 

Department  of  Biology 

North  Carolina  A&T  State  University 

Greensboro,  NC  27411 


ABSTRACT 
On  5  July  1983  a  quantitative  collection  of  bivalves  was  made  at  the  Tangipahoa 
River  in  southern  Mississippi.  Although  the  river  water  was  extremely  soft  and 
the  substrate  and  water  quality  were  good,  a  diverse  and  healthy  mollv^k  com- 
munity was  found.  In  addition  to  the  Asian  Clam.  Corbicula,  five  species  of 
unionids  fFusconaia  flava,  Quadrula  pustulosa,  Elliptic  crassidens,  Tritogonia 
verrucosa  and  Villosa  sp.J  were  taken.  Numerically,  Corbicula  (8.93/m.^),  out- 
numbered total  unionids  (1.2Jt/m.').  However  the  dominant  feature  of  the  benthic 
biomass  were  the  unionids  with  a  total  tissue  dry  mass  of  170^.5  m.g/m'  as  com- 
pared with  1015.0  mglm}  for  Corbicula. 

habitats.  The  purpose  of  this  research  was  to 
characterize,  using  quantitative  field  and 
laboratory  techniques,  a  bivalve  community  in 
the  Tangipahoa  River  in  southern  Mississippi. 


Mississippi  bivalves  were  first  investigated  by 
Hinkley  (1906),  later  by  Grantham  (1969),  and 
then  Stern  (1976)  who  concentrated  on  the 
unionids  of  the  Lake  Maurepas-Pontchartrain- 
Borgene  drainage  system.  Recent  mollusk 
studies  in  Mississippi  include  an  investigation 
of  mussels  in  the  Big  Black  River  (Hartfield 
and  Rummel  1984),  distribution  of  Corbicula 
(Hartfield  and  Cooper  1983),  and  a  discussion  of 
diversity  and  abundance  of  mussels  in  the  south- 
west portion  of  the  state  (Hartfield  and  Ebert 
1984).  The  above-cited  studies,  while  providing 
useful  information  on  species  composition  and 
ecology,  have  been  directed  toward  qualitative 
assessments  of  the  molluscan  fauna.  Quantita- 
tive unionid  studies,  either  in  Mississippi  or 
other  parts  of  the  country,  are  relatively  uncom- 
mon. The  primary  reason  for  this  is  the  diffi- 
culty of  collecting  significant  numbers  of  live 
mussels  from  a  measured  area.  Benthic  grab 
samplers  usually  do  not  retrieve  enough  sub- 
strate to  provide  reliable  population  estimates; 
in  addition,  they  do  not  function  well  in  gravel 


Study  Area 

The  Tangipahoa  River  originates  in  Lincoln 
County,  flows  south  through  Amite  and  Pike 
counties,  then  enters  Lake  Tangipahoa  approxi- 
mately 4  miles  southeast  of  McComb  in  southern 
Mississippi.  Below  Lake  Tangipahoa  the  river 
flows  in  an  easterly  direction  into  Louisiana  and 
ultimately  enters  Lake  Pontchartrain  near  New 
Orleans.  The  study  site  (latitude  31°07'15"N., 
longitude  90°29'25"W.,  elevation  85.3  M.  Mc- 
Comb S,  Miss.,  US  Geological  Survey  Quad- 
rangle) was  located  south  of  Lake  Tangipahoa 
and  2  miles  south  of  the  town  of  Magnolia.  In 
this  area  the  river  consisted  of  pools,  riffles,  and 
runs  with  steep,  partially  eroding  banks. 
Canopy  cover  was  usually  complete  and  sur- 
rounding land  was  either  undeveloped  riparian 
forest  or  pasture.  At  the  time  of  collection  the 


Vol.  100(1) 


January  31, 1986 


THE  NAUTILUS       19 


water  depth  was  no  greater  than  1.0  m,  except 
in  pools,  and  widths  ranged  from  4  to  10  m. 
Based  upon  preliminary  surveys,  this  site  was 
judged  to  be  one  of  the  richest  for  bivalves  when 
compared  with  adjacent  rivers  or  other  portions 
of  the  Tangipahoa  River. 

Substrate  at  the  study  site  consisted  of  gravel- 
ly sands.  On  5  July  1983  chemical  conditions  of 
the  river  water  were  as  follows:  calcium  hard- 
ness, 3.2  mg/1;  pH,  6.8;  and  turbidity,  8.9  NTU. 
On  12  August  at  1300  hr  the  air  temperature 
was  27°C  and  water  temperature  was  24°C. 
Additional  chemical  data  from  the  Tangipahoa 
River  at  Hwy  190  in  Louisiana  (taken  from  the 
Louisiana  Stream  Control  Commission  as  cited 
by  Stern  1976)  further  characterize  this  river  as 
clear,  well  oxygenated,  and  with  low  dissolved 
solids  and  turbidity. 

Methods  and  Materials 

Mollusks  were  collected  from  two  adjacent 
riffles  by  two  workers  during  a  6-hr  period  on  5 
July  1983.  Unionids  were  obtained  from  both 
sites  (214  m^)  and  Corbicula  were  taken  only 
from  a  subsection  of  the  downriver  site  (44  m^). 
Bivalves  were  the  dominant  feature  of  the  com- 
munity; no  snails  or  fingernail  clams  and  only  a 
few  immature  insects  were  collected.  Collec- 
tions were  made  by  hand  or  with  a  basket 
dredge,  an  open-ended  wire  basket  that  was 
pulled  through  the  substrate  and  retained  only 
objects  equal  to  or  greater  than  1  cm  in  dia- 
meter. All  mollusks  were  kept  cool  and  returned 
to  the  laboratory  alive.  The  majority  of  the 
specimens  were  used  in  a  series  of  laboratory 
experiments  on  navigation  effects  that  were 
part  of  the  Environmental  and  Water  Quality 


Operational  Studies  at  the  Waterways  Experi- 
ment Station. 

In  the  laboratory,  total  shell  length  and  height 
were  recorded.  Anterior  and  posterior  mussels 
were  cut  with  a  scalpel  or  small  knife  and  the 
viscera  removed.  Shells  and  viscera  were  dried 
for  24-48  hr  at  75  °C  before  weighing  to  the 
nearest  0.1  mg  on  a  Mettler  balance.  Regression 
equations  for  length,  height,  or  tissue  dry  mass 
(TDM)  were  derived  after  transforming  data  to 
logio  or  logj  with  an  IBM  4331  computer  using 
the  Statistical  Analysis  System. 

Results  and  Discussion 

At  the  study  area  on  the  Tangipahoa  River, 
Corbicula,  which  was  common  in  Mississippi  by 
the  late  1960's  (Grantham  1969),  represented 
87.8%  of  the  total  bivalve  fauna  (Table  1).  Asian 
Clams  were  common  throughout  the  downriver 
site,  even  in  shallow  water  on  unanchored  sands 
where  unionids  were  never  collected.  Length- 
frequency  analysis  (Fig.  1)  indicated  that  two 
major  cohorts  were  present  in  early  July.  About 
40%  of  the  Corbicula  were  approximately 
14-mm  long;  the  second  cohort  ranged  from  20 
to  30  mm.  Based  on  Corbicula  collections  made 
in  May  in  the  Altamaha  River,  Georgia,  Sickle 
(1979)  identified  a  cohort  at  14  mm  and  a  second 
at  22  mm.  In  that  river  the  first  year  spawn  had 
achieved  4  mm  by  August  of  the  same  year.  In 
the  Tangipahoa  River  the  second  year  cohort 
was  more  slow  growing  and  exhibited  a  wider 
range  of  lengths  than  did  the  first  year  group, 
findings  which  are  consistent  with  Sickle  (1979). 
A  power  curve  (Fig.  2)  provided  the  highest  r^ 
value  for  the  relationship  between  shell  length 
and  TDM.  Sickle  (1979)  concluded  that  the  rela- 


TABLE  1.  Summary 

statistics  for  bivalves  collected  from  two 

adjacent  sites  on 

the  Tangipahoa  River 

Pil^eC 

3unty, 

Mississippi,  5  July  1983. 

Total 

Numbers 

Tissue 
TDM/m^ 
1015.0 

Dry 

Mass 

% 
37.3 

Shell    Len 

gth,    mm 

Species 

No. 

Collected 
393 

No./m 
8.93 

% 
87.8 

X 
21.6 

CV* 
23.6 

min 
10.0 

max 

Corbiau la 

33.0 

Fueconaia  flava 

158 

0.74 

7.3 

531.1 

19.5 

46.0 

16.1 

31.3 

70.0 

Quadrula  pustulosa 

55 

0.26 

2.5 

296.8 

10.9 

47.6 

11.8 

36.6 

64.2 

Eltiptio  crassidens 

31 

0.  U 

1.4 

605.5 

22.3 

105.9 

10.9 

84.0 

126.0 

Tritogonia  vevruoosa 

18 

0.08 

0.8 

271.1 

10.0 

94.0 

13.1 

69.0 

108.0 

Villoea  sp. 

4 

0.02 

0.2 

- 

- 

52.9 

15.3 

43.8 

61.1 

Total  unionids 

266 

1.24 

12.2 

1704.5 

62.7 

Total   bivalves 

659 

10.  17 

2719.5 

Coefficient   of  variation 


20      THE  NAUTILUS 


January  81,  1986 


Vol.  100(1) 


CORBICULA 


Lr 


15  20 

SHELL  LENGTH.  MM 


FK;.  1.  Shell  length-frequency  distributions  for  Corhiaila 
collected  from  the  Tangipahoa  River.  Pike  County. 
Mississippi. 


400 
300 

2001- 


100 
80 
60 

40 


CO 


20 


GO 

S2     10 


CORBICULA 
Y  =  0.0047  X3  21  r2  =  0.97 


_L 





J L 


J 


12  5  10 

SHELL  LENGTH.  MIVI 


15    20      30    40 


FIG.  2.  Relationship  between  tissue  dry  mass  and  shell 
length  for  CorbicuUi  collected  from  the  Tangipahoa  River. 
Pike  County,  Mississippi. 


>- 

CJ 


a 

LU 

az 


30   r 
25 

20    h 
15 
10 


o 
ac 


>■ 
o 


a 


o 
cc 


o 


CJ 

cc 


>■ 
o 


¥- 


A 


QUADRULA 
PUSTULOSA 


tfk 


0  20    30    40     50     60     70 
SHELL  LENGTH,  MM 
17 
15   h 


10 


0 


^ 


FUSCONAIA 
FLAVA 


EL 


0  30    40     50     60     70    80 
SHELL  LENGTH,  MM 

25   r 
20 

15   h 
10 
5 

0  Uj 1    III    I" 


TRITOGONIA 
VERRUCOSA 


0  50    60    70     80     90    100  110 
SHELL  LENGTH,  MM 
15  r 


10 


ELLIPTIO 
CRASSIDENS 


0  80     90    100   110   120  130 
SHELL  LENGTH,  MM 

FIC.  3.  Shell  length-frequency  distributions  for  four  species 
of  mussels  collected  from  the  Tangi[)ahoa  River.  Pike 
County,  Mississippi. 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      21 


tionship  between  length  and  TDM  was  best 
described  with  a  power  curve  for  Altamaha 
River  fauna  and  collections  made  in  the  Mud 
River,  West  Virginia,  by  Joy  and  McCoy  (1975). 
Although  Corhicula  numerically  dominated  the 
Tangipahoa  River  bivalve  community,  because 
of  their  smaller  size  they  constituted  only  37.3% 
of  the  total  molluscan  TDM  (Table  1). 

Ficsconaia  Jlava' ,  a  widely  distributed  unionid 
in  southern  Mississippi  and  Louisiana  (Stern 
1976),  was  the  most  abundant  unionid  in  the 
Tangipahoa  River  with  densities  equal  to 
0.74/m^  The  second  most  abundant  species, 
Quadrula  pustulosa.  (this  southern  form  called 
refidgens  Lea  by  some)  has  been  collected  in 
mud,  sand,  and  gravel  substrate  in  flowing  and 
slack  water  habitats  in  this  region  (Stern  1976). 
Total  shell  length  for  F.  Jlava  ranged  from  31.3 
to  70.0  mm;  length-frequency  distributions  (Fig. 
3)  suggest  that  a  major  cohort  existed  from  36 
to  44  mm.  For  Q.  pustulosa  a  cohort  was  identi- 
fied at  approximately  45  mm  with  at  least  one 
below  and  possibly  two  cohorts  above  this 
range.  For  both  of  these  Tangipahoa  River 
unionids,  the  highest  r'  for  length  and  TDM  was 


'Alias  cerina  Conrad.   1838.   See  Hartfield  and   Rummel 
(198.5). 


a  function  of  X'  (Fig.  4).  Quadrula  pustulosa  ex- 
hibited a  slightly  higher  ratio  of  TDM  to  shell 
length,  an  indication  of  greater  tissue  mass  com- 
pared to  length  of  shell,  than  did  F.  flava.  While 
F.  JJava  and  Q.  pustulosa  comprised  only  7.3% 
and  2.5%  of  the  bivalve  community,  because  of 
their  larger  size  they  represented  19.5%  and 
10.9%  of  the  total  bivalve  TDM,  respectively. 

While  Elliptio  crassidens  and  Tritogonia  ver- 
rucosa exhibited  densities  of  0.14  and  0.08/m^ 
respectively,  the  contribution  of  these  larger 
bivalves  to  the  total  TDM  was  approximately 
equal  to  that  of  the  previous  two  species  (22.3% 
and  10.0%,  respectively).  Grantham  (1969) 
reported  that  the  former  species  had  sporadic 
distribution  in  Mississippi,  and  Stern  (1979) 
found  E.  crassidens  common  in  headwater 
streams  in  this  region.  Average  shell  length  of 
E.  crassidens  (105.9  mm)  exceeded  that  for  T. 
verrucosa  (94.0  mm)  by  approximately  10%.  A 
linear  plot  of  total  shell  length  to  total  shell 
height  (Fig.  5)  demonstrated  a  greater  ratio  of 
height  to  length  for  E.  crassidens  then  T.  verru- 
cosa at  sizes  greater  than  97  mm.  For  individ- 
uals larger  than  97  mm,  the  reverse  was  true; 
height  to  length  was  greater  for  T.  verrucosa 
than  for  E.  crassidens. 

In  this  section  of  the  Tangipahoa  River,  F. 


2250 

2000 

1750 

1500 

1250 

1000 

750 

500 

250 


QUADRULA  PUSTULOSA 

Y  =  0.68  X2  -  411.4,  r2  =  0.86 


FISCONAIA  FLAVA 
••    ,,^.,  •.:  Y  =  0.52  X2  -  416.1.  r2  =  0.82 


^,l-J I I I I I I         I 


_L 


0      32     34     36      38     40     42     44     46     48     50      52     54     56     58     60     62     64     66     68 

SHELL  LENGTH.  MM 

FIG.  4.  Relationship  between  tissue  dry  mass  and  shell  length  for  Q.  fnistulosa  and  F.Jhim  collected 
from  the  Tangipahoa  River,  Pike  County,  Mississippi. 


22      THE  NAUTILUS 
80  - 

70  - 

s    60  - 


S    50 


OD     40 


30 


20 


January  81,  1986 

ELUPTIO  CRASSIDENS 

Y  =  0.46X  +  12.3.  r2  =  0.76 


Vol.  100(1) 


TRITOGONIA  VERRUCOSA 
Y  =  0.68X  -  11.7.  r2  =  0.90 


h^ 


_L 


_L 


_L 


_L 


_L 


50 


60 


110 


120 


130 


70    80     90    100 

SHELL  LENGTH.  MIVI 

FIG.  .5.  Relationship  between  shell  height  and  shell  length  for  E.  crassideiis  and  T.  verrucosa  col 
lected  from  Tangipahoa  River,  Pike  County,  Mississippi. 


flava,  because  of  its  numbers,  and  E.  crassidens. 
because  of  its  size,  dominated  the  unionid  TDM. 
The  other  three  unionids,  Q.  pustulosa,  T.  verru- 
cosa, and  Villosa  sp.,  were  a  minor  part  of  the 
unionid  community.  While  the  unionids  domi- 
nated the  bivalve  biomass  (62.7%),  they  were 
outnumbered  by  Corbicula  (87.8%).  In  addition, 
the  Asian  clams,  because  of  their  rapid  growth 
rates,  exhibited  higher  productivity  in  terms  of 
tissue  growth  per  unit  time  than  did  the 
unionids. 

Since  dissolved  calcium  is  of  obvious  impor- 
tance for  shell  maintenance,  it  is  significant  that 
the  Tangipahoa  River  supported  a  healthy  com- 
munity regardless  of  soft  water  (3.2mg/(  as 
CaCOj).  In  central  New  York,  Clarke  and  Berg 
(19.59)  collected  no  unionids  in  water  with  total 
alkalinity  less  than  47  ppm.  Harman  (1969) 
found  unionids  in  water  with  total  alkalinity 
ranging  between  20  and  30  mg/f  and  stated  that 
sudden  changes  in  pH  were  probably  more  detri- 
mental than  low  dissolved  solids.  Tangipahoa 
River  fauna  live  in  water  at  the  extreme  lower 
end  of  the  range  of  dissolved  calcium  concentra- 
tions usually  reported  for  the  freshwater 
Mollusca. 

A  site  on  the  Tangipahoa  River  in  southern 
Mississippi   was  identified   which  supported  a 


dense,  fairly  diverse  mollusk  community. 
Bivalves  were  found  in  good  quality  substrate  in 
water  that  was  clear,  well  oxygenated  but  with 
extremely  low  dissolved  calcium  levels.  While 
freshwater  mollusks  are  usually  found  in 
medium  to  hardwater  habitats,  it  appears  that 
reduced  dissolved  calcium  concentrations  are 
not  limiting  the  Tangipahoa  River  fauna. 

LITERATURE  CITED 

Clarke.  A.  H.,  Jr.  and  Berg,  C.  0,  19.59.  The  Freshwater 
Mussels  of  Central  New  York,  with  an  Illustrated  Key  to 
the  Species  of  Northeastern  North  America.  University 
Agricultural  Experiment  Station  Memoir  367:1-79. 

Grantham,  B.  J.  1969.  The  Freshwater  Pelecypod  Fauna  of 
Mississippi,  Doctoral  Dissertation,  University  of  Southern 
Mississippi,  Hattiesburg,  Miss. 

Harman.  W.  N.  1969.  The  Effect  of  Changing  pH  on  the 
Unionidae.  The  Nautilus  83:69-70. 

Hartfield,  P.  D.  and  Cooper,  C.  M.  1983.  Distribution  of 
Corbicula  Jluminea  the  Asiatic  Clam,  in  Mississippi.  The 
Nautilus  97:66-68. 

Hartfield,  P.  D.  and  Ebert,  D.  1984.  Factors  Affecting  the 
Diversity  and  Abundance  of  Mussels  in  Southwest  Missis- 
sippi. Presentation  for  Mississippi  Academy  of  Sciences, 
Biloxi,  Miss.,  March  1984. 

Hartfield,  P.  D.  and  Rummel,  R.  G.  198,5.  Freshwater 
Mussels  (Unionidae)  of  the  Big  Black  River,  Mississippi. 
The  Nautilus  99(4):\\6-\l9. 

Hiiikley,  A.  A.  1906.  Some  Shells  from  Mississippi  and 
Alabama.  The  Nautilus  20:52-55. 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      23 


Joy,  J.  E.  and  McCoy,  L.  E.  1975.  Comparison  of  Shell 
Dimensions  and  Viscera  Mass  Weights  in  Corbicula 
manilensis  (Philippi,  1844).  The  Nautilus  89(2):51-54. 

Sickle,  J.  B.  1979.  Population  Dynamics  of  Corbicula  in  the 
Altamaha  River,  Georgia.  In  J.  C.  Britton,  ed.,  Proceed- 
ings.   First   International   Corbicula   Symposium,    The 


Texas  Christian  University  Research  Foundation,  Ft. 
Worth,  Texas. 
Stern,  E.  M.  1976.  The  Freshwater  Mussels  (Unionidae)  of 
the  Lake  Maurepas-Pontchartrain-Borgne  Drainage  Sys- 
tem, Louisiana  and  Mississippi.  Ph.D.  Thesis,  Louisiana 
State  University,  Baton  Rouge,  La. 


NOTES  ON  SPECIES  OF  BROCCHINIA 
(GASTROPODA:  CANCELLARIIDAE) 


Richard  E.  Petit' 

Research  Associate 
Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 

Smithsonian  Institution 

Washington,  D.C.  20560 

ABSTRACT 
Available  replacement  names  are  discussed  for  Voluta  mitraeformis  Brocchi, 
181Jt,  the  type  species  of  Brocchinia  Jousseaume,  1887,  and  a  primary  junior 
homonym  of  Voluta  mitraeformis  Lamarck,  1811.  Cancellaria  pusilla  Adams, 
1869  is  also  preoccupied.  This  Recent  species  is  redescribed  as  Brocchinia  clenchi, 
n.  sp. 


Voluta  mitraeformis  Brocchi,  1814,  a  well- 
known  cancellariid  of  the  European  Tertiary, 
is  the  type  species  of  the  genus  Brocchinia 
Jousseaume,  1887.  Although  this  taxon  is  fre- 
quently cited,  its  status  as  a  junior  primary 
homonym  of  Voluta  mitraeformis  Lamarck, 
1811  has  been  noted  in  the  literature  only  by 
Malatesta  (1974:376)  and  DavoH  (1982:61). 

Malatesta  (1974:377)  considered  Cancellaria 
pusilla  H.  Adams,  1869  to  be  available  as  a 
replacement  name  for  the  preoccupied  Voluta 
mitraeformis  Brocchi,  although  he  considered 
the  Recent  and  Tertiary  populations  to  be  sepa- 
rable. He  further  stated  that  the  "fossil  sub- 
species" {sotfospecie  fossile)  had  as  an  available 
name  Cancellaria  cerithiopsis  Almera  &  Bofill, 
1887  {sic).  Malatesta's  conclusions  are  incorrect 
as  C.  pusilla  H.  Adams  is  not  an  available  name, 
being  a  junior  primary  homonym  of  Cancellaria 
pusilla    Sowerby,    1832,    and    C.    cerithiopsis 


'Mailing  address:  P.O.  Box  30,  North  Myrtle  Beach,  SC 
29582. 


Almera  &  Bofill  dates  from  1898,  not  1887. 
Earlier  synonyms  of  V.  mitraeformis  Brocchi 
exist,  as  shown  below. 

Davoli  (1982:61)  stated  that  Voluta  mitrae- 
formis Brocchi,  1814  and  Voluta  mitraeformis 
Lamarck,  1811  should  not  be  regarded  as  pri- 
mary homonyms  inasmuch  as  Brocchi  pointed 
out  that  his  species  belonged  to  Lamarck's 
genus  Cancellaria.  Davoli  referred  to  the 
Preamble  of  the  International  Code  of  Zoologi- 
cal Nomenclature  in  his  plea  for  retention  of 
Brocchi's  V.  mitraefoifmis,  but  the  applicable 
portion  of  the  Code  in  this  instance  is  Article 
60a  which  states:  "A  junior  homonym  must  be 
rejected  .  .  ."  (italics  added).  This  is  unequivocal. 

Brocchinia  Jousseaume,  1887 
Type,  by  subsequent  designation  of  Sacco,  1894,  Brocchi- 
nia mitraeformis  (Brocchi)  ( =  Voluta  mitraeformis  Brocchi, 
1814  non  Voluta  mitraeformis  Lamarck,  1811;  =  Brocchinia 
parvula  tauropama  Sacco,  1894). 

Brocchinia  has  been  cited  as  a  subgenus  of 
Narona  H.  &  A.  Adams,  1854  by  various 
authors  (see  synonymy  in  Malatesta,  1974:377). 


24      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


The  type  of  Narona  is  Cancellaria  clavaiula 
Sowerby,  1832  from  the  tropical  eastern  Pacific. 
There  is  no  evidence  for  a  subgeneric  relation- 
ship between  the  two  genus-level  taxa.  Narona 
is  characterized  by  a  somewhat  tabulate  shell 
having  a  distinct  anterior  canal;  its  columella 
bears  two  strong,  sharp  folds,  the  posterior  one 
being  superior.  Brocchinia  has  a  small  non- 
tabulate  shell  with  a  rounded  anterior;  the  short 
siphonal  canal  evidenced  only  as  a  slight  curving 
of  the  columella;  its  two  columellar  folds  are 
weak  and  rounded. 

Brocchinia  parvula  parinila  (Beyrich,  1856) 

Synoyiymy- 
Cancdlaria  parvula  Beyrich,  18.56:58,  pi.  3,  figs.  82a-b. 
Cancellaria  bicarinata  Homes  &  Auinger,  1890:281,  pi.  33, 

figs.  16a-c. 
Brocchinia    mitraefnrmis    parvula    (Beyrich),    .Jansseii, 

1983:9,  pi.  1,  figs.' 5-7;  pi.  5,  fig.  3,  text-fig.  4. 

Cancellaria  parvula  Beyrich  is  apparently 
the  earliest  named  taxon  considered  to  be  a 
synonym  or  subspecies  of  Valuta  vfiitraeformis 
Brocchi,  and  therefore  becomes  the  nominotypi- 
cal  subspecies  of  the  species  group  (I. C.Z.N. 
Article  47).  This  German  Miocene  species  was 
cited  by  Gilbert  (1952:130,  pi.  9,  fig.  16;  1960:4) 
as  a  form  of  C.  mitraeformis  (Brocchi),  his 
figure  agreeing  well  with  Beyrich's  original 
figures.  Brocchinia  parvula  has  been  treated  as 
a  valid  species,  separable  from  B.  mitraeformis. 
by  Kautsky  (1925:137),  Sacco  (1894:70)  and 
Seiber  (1936:93).  Davoli  (1982:61)  mentions  C. 
parvula  in  his  discussion  but  does  not  include  it 
in  his  synonymy  of  B.  mitraeformis.  Malatesta 
(1974:376),  in  his  synonymy,  lists  Kautsky's  cita- 
tion of  B.  panmla  but  does  not  list  the  original 
description,  a  possible  indication  that  he  consid- 
ered the  two  to  be  separable.  In  the  most  recent 
paper  treating  the  subject,  Janssen  (1983:9,  10) 
considers  B.  parvula  to  be  a  subspecies  of.S. 
mitraeformis  (Brocchi)  and  states  that  B.  par- 
vula occurs  in  the  Mediterranean  Pliocene  as 
well  as  in  the  northern  European  Miocene.  This 
interpretation  of  the  relationship  between  B. 
parvula  and  B.  mitraeformis  agrees  with  this 
writer's  opinion. 

Brocchinia  parvula  tauroparva  Sacco,  1894 
Synonymy  - 
Volula  milraiiformis  Brocchi,  1814:fi45,  pi.  15,  fig.  13  (noii 
Valuta  mitraeformis  Lamarck,  1811:73). 


Brocchinia  ynitraeformix  var.  tauroparva  Sacco,  1894:68, 

pi.  3,  fig.  82. 
Narona  (Brocchinia)  mitraejhi-viis  (Brocchi),  Davoli,  1982: 

61,  pi.  7,  figs.  3-6. 

Sacco  (1894:68-70)  named  eight  varieties 
of  Brocchinia  mitraeformis  (Brocchi)  from 
the  Italian  Pliocene.  The  first  of  these,  Broc- 
chinia mitraeformis  tauroparva.  is  not  distinct 
from  the  typical  form,  and  is  here  considered  to 
be  a  subjective  synonym  of  B.  mitraeformis 
(Brocchi).  Brocchinia  tauroparva  (Sacco)  thus 
replaces  the  preoccupied  V.  mitraeformis 
Brocchi. 

Brocchinia  clenchi,  n.  sp. 

'  Figs.  1-4 

Synonymy - 

Cancellaria  pufiiJla  H.  Adams,  1869:274,  pi.  19,  fig.  ]2(non 
Cancellaria  puxilla  Sowerby.  1832:6,  fig.  34). 

Cancellaria  mitraeformis  Brocchi,  .Jeffreys,  1885:49. 

Narona  (Brocchinia)  p^isilla  (H.  Adams),  Nordsieck,  1968: 
151,  pi.  25,  fig.  87.,3(). 

Df'.srnj9^iOK-Paucispiral  nucleus  smooth, 
heliciform,  consisting  of  about  IV2  whorls;  tran- 
sition from  protoconch  to  teleoconch  almost  im- 
perceptible. Faint  peripheral  spiral  cords  arise 
midway  on  the  first  postnuclear  whorl.  The 
spiral  sculpture  consists  of  3  to  5  extremely 
weak  cords,  with  3  usually  visible  on  the  spire 
whorls,  and  5  or  so  on  the  body  whorl.  Axial 
sculpture,  when  present,  consists  only  of  weak 
nodes  on  the  periphery  that  are  crossed  by  the 
spiral  cords.  Suture  distinct,  slightly  impressed. 
Teleoconch  of  about  4  whorls;  nonumbilicate. 
Aperture  ovate  with  a  weak  but  noticeable  col- 
umellar callus.  Columella  with  two  distinct, 
rounded  folds  of  approximately  equal  size,  the 
posterior  one  being  slightly  larger  and  more 
pronounced.  Siphonal  canal  indicated  by  the 
angled  base  of  the  columella  and  a  minute 
depression  in  the  base  of  the  aperture.  Outer  lip 
prosocline,  usually  denticulate  within.  The  den- 
ticles, which  do  not  extend  to  the  edge  of  the  lip, 
vary  in  strength  and  number.  Shell  white  or 
horn  colored.  Animal  unknown. 

Type  Material:  Holotype,  4.5  mm  x  2.3  mm 
(USNM  849002);  Paratype,  5.2  mm  x  2.6  mm 
(USNM  189694);  Paratype,  BM(NH)  1855.4. 
4.202,  6.0  mm  x  2.5  mm  (possibly  the  type  of  C. 
pu.'iilla  Adams). 

Type  locality:  Josephine  Bank,  340-430  fms. 
Josephine  Bank  is  situated  at  approximately 


Vol.  100(1) 


January  31, 1986 


THE  NAUTILUS      25 


FIGS.  1-4.  Brricrhima  rlenchi  new  species.  1,  Paratype,  BM(NH)  185.S.4.4.202,  Oratova,  Canary 
Islands.  15x.  2.  Paratype,  USNM  189694,  Josephine  Bank,  621-786  m,  20x.  3,  Holotype,  USNM 
849002,  Josephine  Bank,  621-786  m,  20  x.  4,  Apical  view  of  protoconch  of  holotype,  65  x. 


37°N,  14°W,  due  West  of  the  southern  tip  of 
Portugal.  The  type  and  paratype  in  the  USNM 
are  from  the  Jeffrey's  collection. 

Etymology:  This  species  is  respectfully  dedi- 
cated to  the  memory  of  the  late  Dr.  William  J. 
Clench. 

Discussion:  This  Recent  species  was  previous- 
ly described  as  Cancellaria  pusilla  Adams, 
1869,  but  that  name  is  preoccupied  by  C.  pusilla 
Sowerby,    1832.   Jeffreys  (1885:49)  placed  C. 


pusilla  Adams  in  the  synonymy  of  C.  rnitrae- 
formis  (Brocchi),  while  Sykes  (1911:332), 
Harmer  (1918:396)  and  Dautzenberg  (1927:73) 
considered  the  Recent  species  to  be  distinct. 

The  Recent  B.  clenchi  is  easily  separable  from 
the  Tertiary  species  by  its  much  smaller  size  and 
by  its  comparatively  weak  sculpture.  The  illus- 
tration accompanying  Adams'  description  of  C. 
pusilla  is  misleading,  as  the  fine,  almost  in- 
distinct spirals  are  shown  to  be  quite  strong. 


26      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


The  drawings  of  Nordsieck  (1968:pl.  25,  fig. 
87.30;  1979:pl.  37,  fig.  6)  are  too  poor  to  be 
recognizable.  Although  included  by  Nordsieck  & 
Talavera  (1979:152)  with  the  notation  that  its 
occurrence  in  the  Canaries  may  be  accidental, 
specimens  have  now  been  found  in  fish  traps  in 
Tenerife  South  at  a  depth  of  43  m  (Talavera, 
pers.  comm.). 

As  the  type  of  C.  piisilla  Adams  cannot  be 
positively  identified,  the  species  has  been  re- 
described  so  that  a  holotype  can  be  designated, 
rather  than  simply  proposing  a  nomen  novum. 
The  BM(NH)  specimen  listed  as  a  paratype  is 
from  the  R.  M.  McAndrew  collection. 

Acknowledgments 

This  paper  would  not  have  been  possible  with- 
out considerable  assistance.  Dr.  Franco  Davoli, 
Universita  di  Modena,  Italy,  furnished  a  transla- 
tion of  his  work,  and  corresponded  regarding 
the  desirability  of  retaining  Brocchi's  taxon.  Mr. 
Piero  Piani,  Bologna,  Italy,  furnished  transla- 
tions, comments  and  literature. 

Mrs.  Kathie  Way,  British  Museum  (Natural 
History),  London,  located  the  possible  type  of 
Adams'  C.  pusilla  and  made  it  available  for 
study. 

The  late  Dr.  Joseph  Rosewater,  National 
Museum  of  Natural  History,  Washington,  D.C., 
made  the  Jeffreys  specimens  available  and 
located  pertinent  references. 

Dr.  Francisco  G.  Talavera,  Museo  Insular  de 
Ciencias,  Tenerife,  Canary  Islands,  furnished  in- 
formation and  literature. 

Dr.  A.  G.  Beu,  N.Z.  Geological  Survey,  Lower 
Hutt,  New  Zealand,  read  and  commented  on  an 
early  draft  of  this  paper. 

Dr.  M.  G.  Harasewych,  National  Museum  of 
Natural  History,  Washington,  D.C.  made  the 
photographs,  prepared  the  plate,  and  reviewed 
the  manuscript. 

The  cooperation  and  assistance  of  all  is  grate- 
fully acknowledged. 


LITERATURE  CITED 

Adams.  H.  1869.  Descriptions  of  a  new  genus  and  fourteen 

new  species  of  marine  shells.  Froc.  Zool.  Soc.  Loiidtni. 

1869^112-21^,  pi.  19. 
Adams,  H.  &  A.  1853-1858.  The  genera  uf  Recent  Mollusca. 

3  vols.  London. 
Almera,  J.  &  A.  Bofill.  1898.  Moluscos  fosiles  recogidos  en 

los  terrenos  pliocenicos  de  Cataluna.  Bol.   Com.  Mapa 


geoL  Espana.  IV  (ser.  2):  XII  +  223,  14  pis.,  Madrid. 
Beyrich,  E.  1856.  Die  Konchylien  des  norddeutschen  Ter- 

tiargebirges.  Zeitschr.  d.  Deutsch.  geol.  Ges.  8. 
Brocchi,  G.  B.  1814.  Conchiologia  fossile  subappenniria  con 

osservazioni    geotogirhe    sugli    Appenniyii.     2     vols., 

Stamperia  Reale,  Milano. 
Dautzenberg,   Ph.    1927.   Mollusques  provenant  des  cam- 

pagnes  scientifiques  du  Prince  Ibert  I  de  Monaco  dans 

rOcean    Atlantique   et   le    Golfe   de   Gascogne.   Result. 

Camp.  Scient.  Pr.  Alberta  I  de  Monaco,   fasc.  LXXII, 

400  pp.,  9  pis. 
Davoli,  F.  1982.  Cancellariidae  (Gastropoda).  In  E.  Monta- 

naro  Gallitelli  (ed.).  Studi  monografici  sulla  malacologia 

mioceniea  modenese.  Parte  I  -  I  Molluschi  tortoniani  di 

Montegibbio.  Pobieont.  Ital  72  (n.ser.  vol.  42);5-73,  7  pis., 

Pisa. 
Glibert,  M.  1952.  Faune  malacologique  du  Miocene  de  la 

Belgique.  II.  Gastropods.  Inst.  Roy.  Sci.  Nat.  Belg.,  Mem. 

121:1-197,  10  pis. 
1960.  Gastropodes  du  Diestien.  du  Scaldisien  et 

du  Merxemien  de  la  Belgique.  4me  Note.  Inst.  Roy.  Sci. 

Nat.  Belg..  Bull.  36(33):  1-44,  2  pis. 
Harmer,  F.  W.  1918.  The  Pliocene  Mollusca  of  Great  Britain. 

Part  III.  Pnheontographical  Soc.  70:303-461.  pis.  33-44. 
Horncs,   R.   &   M.   Auinger.    1890.   Die  Gastropoden  der 

Meeres-Ablagerungen  der  ersten  und  zweiten  Miocanen 

Mediterranstufe  in  der  ostereich-ungarischen  Monarchie. 

Abhdlg.  Geol.  Reichsanst.  6. 
International    Commission    on    Zoological    Nomenclature. 

1985.    International   Code  of  Zoological   Nomenclature, 

3rd  Ed.  pp.  i-xx,  1-338.  London. 
Janssen,  A.  W.   1983.   An  account  of  the  Cancellariidae 

(Gastropoda)  of  Winterswijk-Miste  (Miocene.  Hemoorian), 

The  Netherlands.  Scripta  Geol.  68:1-39.  6  pis. 
Jeffreys,  J.  G.  1885.  On  the  Mollusca  procured  during  the 

'Lightning'  and  'Porcupine'  Expeditions,   1868-70.  (Part 

IX).  Proc.  Zool.  Soc.  London.  1885:27-63 ,  pis.  4-6.  . 
Jousseaume,  F.  P.  1887.  La  famille  des  Cancellariidae.  Le 

Natural.,   ami.   9.   2e  ser.,   pp.    155-157,    163,    192-194, 

213-214,  221-223. 
Kautsky,  F.  1925.  Das  Miocan  von  Hemmoor  und  Basbeck- 

Osten.  Abh.   preuss.   geol.   LandesanM..  N.F.   97:1-255, 

12  pis. 
Lamarck,   .J.B.P.A.    1811.   Suite  de  la  determination  des 

especes  de  Mollusques  testaces.  Valuta.  Ann.  Mus.  Nat. 

d  Hist.  Nat.  17:54-80. 
Malatesta,  A.  1974.  Malacofauna  pliocenica  umbra.  Mem. 

Carta  Geol.  It  13:XI-i-498,  32  pis..  Roma. 
Nordsieck,  F.  1968.  Die  europaischen  Meeres-Gehausesck- 

necken  (Prosobranrhia)  com  Eisnifer  bis  Kapverden  U7id 

Mittelmeer.  268  p.  Gustav  Fischer.  Stuttgart. 
Nordsieck,  F.  &  F.  G.  Talavera.  1979.  Moluscos  Marinas 

de  Canarias  y  Madera  (Gastropoda).   208   p.,   46  pis. 

Tenerife. 
Sacco,  F.  1894.  I  mollu.schi  dei  terreni  terziarii  del  Piemonte 

e  Liguria.  Pt.  XVI  (Cancellariidae).  Mem.  R.  Accad.  Sc. 

Torino,  Clausen  Ed.,  78  pp..  3  pis..  Torino. 
Sieber,  R.  1936,  Die  Cancellariidae  des  niederosterreichis- 

chen  Mioziins.  Arrhir.  Mollu.fk.  68:65-115. 
Sowerby,  G.   B.   1832-33.  The  Conchological  Illustrations. 

Cancellaria.  Pts.  9-13.  London.  5  pis.  with  explanations 

-I-  catalogue,  10  pp.  (Pts.  9-12,  figs.  1-35,  published  1832; 


Vol.  100  (1) 


January  31, 1986 


THE  NAUTILUS      27 


pt.  13,  figs.  36-44  and  catalogue  published  1833). 
Sykes,  E.  R.  1911.  On  the  mollusca  procured  during  the 


"Porcupine"  Expeditions,  1869-70.  Supplemental  Notes, 
Part  IV.  Proc.  Malac.  Soc.  London  9(6):331-348. 


ON  THE  TYPE  SPECIES  OF  METULA  H.  &  A.  ADAMS,  1853: 

BUCCINUM  CLATHRATUM  A.  ADAMS  AND  REEVE,  1850 

(GASTROPODA:  BUCCINIDAE) 


William  K.  Emerson 

Department  of  Invertebrates 

American  Museum  of  Natural  History 

New  York,  New  York  10024 

ABSTRACT 

The  type  species  o/Metula  H.  and  A.  Adams.  1853:  Buccinum  clathratum  A. 
Adams  and  Reeve,  1850  (not  Kiener,  183J,,  nor  Anton,  1839)  is  determined  to  be 
referable  to  Metula  amosi  Vanatta,  1913,  from,  the  tropical  eastern.  Pacific.  The 
genera  Acamptochetus  Cossmann,  1901;  Antemetula  Rehder,  19J,3;  and  Colubra- 
rina  Kuroda  and  Habe,  in  Kuroda,  Habe,  and  Oyama,  1971,  are  placed  in  the 
synonymy  of  Metula. 


The  "Metula  problem"  has  long  been  the  con- 
cern of  workers  dealing  with  these  buccinid 
gastropods  owing  to  the  uncertain  nomencla- 
tural  and  taxonomic  status  of  the  type  species  of 
this  genus-group  taxon  (E.  A.  Smith,  1904; 
Woodring,  1928;  Tomlin,  1927;  Rehder,  1943; 
Altena,  1949;  Knudsen,  1956;  Cernohorsky, 
1971;  Olsson  and  Bayer,  1972,  Kilburn,  1975; 
and  Houbrick,  1984).  This  note  undertakes  to 
solve  these  questions. 

The  type  of  Metula,  Buccinum.  clathratum 
A.  Adams  and  Reeve  (1850,  p.  32,  pi.  11,  fig. 
12),  was  stated  to  have  been  dredged  off  the 
Cape  of  Good  Hope,  in  136  fathoms  [248  meters] 
during  the  1843-1846  voyage  of  the  H.M.S. 
"Samarang".  TomHn  (1927,  p.  160),  in  his  review 
of  the  South  American  mollusks  of  the  "St. 
George"  Expedition,  however,  noted  that  this 
species  was:  "Originally  described  from  deep 
water  off  the  Cape,  but  that  locality,  like  many 
others  in  the  'Samarang'  work,  is  certainly 
erroneous."  Furthermore,  Tomlin  (1927,  op.  cit.) 
concluded  that  this  taxon  was  referable  to  speci- 
mens in  his  collection  from  Balboa,  Panama, 
which  he  identified  as  "Metula  clathrata  (A.  Ad. 
&  Rve.)".  Because  later  workers  have  largely  ig- 
nored or  questioned  the  identity  of  Buccinum 


clathratum  A.  Adams  and  Reeve,  1850,  the  type 
species  oi  Metula  H.  and  A.  Adams,  I  examined 
the  holotypic  specimen  of  this  taxon,  which  is  in 
the  British  Museum  (Natural  History). 

The  specimen  labeled  as  holotype  of  Buccinum 
clathratum.  (BM(NH),  1874. 12. 1'l.  145;  here  illus- 
trated, figures  1,  2)  is  25.3  mm  in  height.  The 
original  polychrome  illustration  of  the  type  (A. 
Adams  and  Reeve,  1850,  pi.  11,  fig.  12)  depicts  a 
specimen  of  the  same  size  as  the  holotype,  which 
is  now  faded  and  has  a  chip  on  the  anterior  por- 
tion of  the  outer  lip. 

As  Cernohorsky  (1971,  p.  149)  has  pointed 
out,  Bu.cci.num  clathratum  (A.  Adams  &  Reeve, 
1850)  is  twice  preoccupied  (not  Kiener,  1834,  p. 
101,  nor  Anton,  1839,  p.  91).  Fortunately,  a  re- 
placement name  is  not  required,  as  Metula 
amosi  Vanatta  (1913,  p.  22,  figs.  1,  2;  Keen, 
1971,  p.  566,  fig.  1133;  Olsson  and  Bayer,  1972, 
pp.  906,  907,  figs.  1,  B-D)  is  an  available  junior 
synonym  of  this  taxon.  The  most  common  of  the 
Panamic  Metula,  this  species  ranges  offshore 
from  the  Gulf  of  California  to  Panama  Bay. 
Although  large  specimens  attain  more  than  44 
mm  in  height,  mature  examples  of  M.  amosi  in 
the  American  Museum  of  Natural  History 
(AMNH)  collection  from  Mexico  and  Panama 


28      THE  NAUTILUS 


.lanuarvSl,  1986 


Vol.  100  (1) 


are  nearly  as  diminutive  as  the  lioiotype  of  M. 
clathratn.  Tomiin's  (1927,  op.  cit.)  referral  of 
this  species  to  Pacific  Panamanian  waters  is 
thus  confirmed,  and  the  type  locality  of  M. 
amosi  is  here  restricted  to  Balboa,  Bahia  de 
Panama. 

It  should  be  noted  that  Knudsen  (1956,  pp.  39, 
40,  pi.  1,  fig.  1)  referred  a  specimen  dredged  off 
"Spanish  Guinea",  West  Africa  to  Metula 
clatkrata  (A.  Adams  &  Reeve,  1850)  (not 
Kiener,  1834,  nor  Anton,  1839).  The  specimen 
figured  by  Knudsen  is  47  mm  in  height  (here 
reproduced,  fig.  3)  and  is  referable  to  an  uniden- 
tified specimen  of  Metula  in  the  AMNH  collec- 
tion (#198755;  here  illustrated,  figs.  4,  5)  from 
"West  Africa",  cx-Marcel  Pin  collection. 
Another  record  of  this  west  African  species, 
taken  in  250  meters  off  Dakar,  Senegal  (Natal 
Museum  J.  4152)  was  referred  to  me  by  Dr.  R. 
N.  Kilburn.  These  specimens,  however,  are  not 
conspecific  with  Metula  clathrata  (=M.  amosi 
Vanatta)  on  the  basis  of  the  type  specimen  of  A. 
Adams  and  Reeve's  M.  clathrata  (cf .  figures  1 ,  2 
with  figures  3-5),  and  they  appear  to  represent  a 
new  species.  Nor  is  Knudsen's  west  African 
specimen  referable  to  Metula  boswellae  Kilburn 
(1975,  pp.  594,  595,  fig.  10b,  10c),  from  off 
"MoQamhique",  East  Africa.  Kilburn  (1975,  p. 
592)  accepted  Knudsen's  (1956,  op.  cit.)  referral 
of  the  "Spanish  Guinea"  specimen  to  M. 
clathrata  (A.  Adams  &  Reeve,  1850)  and  he 
noted  that  this  taxon  was  preoccupied.  Kilburn 


(1975,  op.  cit.),  therefore,  renamed  this  taxon, 
Metula  knudseni  (Kilburn,  1975,  p.  592).  This 
unnecessary  replacement  name  thus  becomes  a 
junior  subjective  synonym  of  Metula  amosi 
Vanatta,  1913. 

A  number  of  authors  have  discussed  the  taxo- 
nomic  status  oi Metula  H.  and  A.  Adams  (1853, 
p.  84).  Rehder  (1943,  p.  199)  appears  to  be  the 
first  to  attribute  Metula  clathrata  (A.  Adams 
and  Reeve,  1850)  as  the  type  species  (subse- 
quent selection  of  Kobelt,  1876,  p.  29,  pi.  6,  fig. 
11  [  =  "12"]).  Woodring  (1928,  p.  286;  1964,  p. 
259;  Keen,  1971,  p.  566;  Olsson  and  Bayer, 
1972,  p.  902)  considered  Buccinum  m.etula 
Hinds,  1844,  to  be  the  type  species  by  "hidden 
tautonymy,"  for  which  there  is  no  provision  in 
the  ICZN  Code.  Thus,  this  designation  must  be 
rejected  (Cernohorsky,  1971,  p.  149).  Kobelt's 
(1876,  op.  cit.)  selection  oi  Metula  clathrata,  as 
the  type  species,  therefore,  appears  to  be  the 
first  available  designation  for  Metula. 

Cernohorsky  (1971,  pp.  151,  152)  and 
Houbrick  (1984,  p.  420)  considered  Acampto- 
chetus  Cossmann  (1901,  p.  123)  diwd  Antemetula 
Rehder  (1943,  p.  199)  to  be  congeneric  taxa.  The 
type  species  of  Acamptochetus,  by  original 
designation,  is  Murex  mitraeformis  Brocchi, 
1814,  from  the  Neogene  of  Italy  and  is  a  typical 
Metula  (Cernohorsky,  1971,  fig.  46).  Other 
Neogene  fossils  are  known  from  Europe,  Java, 
Sumatra,  India  and  the  Americas  (Altena,  1949; 


FIGS.  1-5.  1  and  2,  Metula  amosi  Vanatta,  holotype  of  Buccinum  clathrnia  A.  Adams  and  Reeve,  BM(NII)  1874.  lli.  I  1 . 1 4;'.. 
x2.  3,  "Metula  clathrata"  after  Knudsen  (19.56,  pi.  1,  fig.  1)  from  "Spanish  Guinea",  not  Metula  clathrata  (A.  Adams  and 
Reeve),  x  IV2.  4  and  5,  Metula  sp.,  AiVINH  1987.5.5,  ex-Marcel  Pin  coll.,  from  "West  Africa",  x  1 V2. 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      29 


Olsson  and  Bayer,  1972;  Malatesta,  1974;  and 
Grecchi,  1978).  The  type  species  oi  Antemetula 
by  original  designation  is  Buccinum  metula 
Hinds,  1844.  This  species  was  originally 
reported  from  the  "West  coast  of  Veragiia 
[Republica  de  Panama] .  .  .  from  a  depth  of  a  few 
fathoms,  among  mud."  This  locality,  however,  is 
apparently  an  error,  as  no  additional  specimens 
have  been  recognized  in  tropical  American 
waters.  The  type  specimen  (Hinds,  1844,  pi.  16, 
figs.  13,  14;  refigured  by  Keen,  1971,  p.  567,  fig. 
1134  and  by  Olsson  and  Bayer,  1972,  fig.  lA)  is 
not  in  the  British  Museum  (Natural  History) 
teste  Kathie  Way,  1985).  Furthermore,  this 
specimen  could  not  be  located  there  some  35 
years  ago  (Altena,  1949,  p.  385).  Many  years  ago 
Smith  (1904,  p.  465)  noted  a  close  resemblance 
of  young  specimens  of  Metula  mitreUa  (A. 
Adams  and  Reeve,  1850,  p.  32,  pi.  11,  fig.  13, 
from  the  "China  Sea;  .  .  .  ten  fathoms"),  to  the 
illustration  of  the  type  specimen  of  M.  metula. 
Smith  (1904,  op.  cit.)  concluded  that  Buccinum 
'metula  was  conspecific  with  B.  mitrella.  a  well- 
known  Indo-West  Pacific  species,  and  that  the 
original  west  American  locality  was  probably 
erroneous.  I  agree  with  his  conclusions  {cf.  the  il- 
lustration of  Cernohorsky  (1971,  p.  150,  fig.  47) 
with  that  of  Olsson  and  Bayer  (1972,  p.  905,  fig. 
lA).  If  this  interpretation  is  correct,  M.  mitrella 
(A.  Adams  and  Reeve,  1850)  is  a  junior  subjec- 
tive synonym  of  M.  metula  (Hinds,  1844)  and 
this  taxon  should  be  removed  from  the  eastern 
Pacific  faunal  list. 

Kilburn  (1975,  p.  592)  assigned  the  genus- 
group  name,  Colubrarina  Kuroda  &  Habe,  in 
Kuroda,  Habe  &  Oyama  (1971,  p.  173)  to  the 
synonymy  of  Metula.  The  type  species  of  this 
taxon  by  original  designation  is  Antemetula 
(Colubrarina)  metulina  Kuroda  &  Habe,  in 
Kuroda,  Habe  &  Oyama,  1971  {op.  cit..  p.  173, 
pi.  46,  fig.  9),  from  Honshu  and  Shikoku,  Japan, 
in  90-200  m.  Described  as  a  subgenus  oiAnteyne- 
tula  Rehder,  the  type  species  of  Colubrarina 
was  said  to  differ  from  the  nominate  subgenus 
by  having  a  larger  and  more  coarsely  sculptured 
shell. 

In  summary,  the  buccinid  genus  Metula  H. 
and  A.  Adams,  1853,  with  the  type  species:  M. 
clathrata  (A.  Adams  and  Reeve,  1850)  [  =  M. 
amosi  Vanatta,  1913  (synonym;  M.  knu(lse7ii 
Kilburn,  1975)],  is  available  for  an  assemblage  of 


Neogene  species  from  the  Old  and  New  World 
tropical  belt  and  for  several  Recent  species  sur- 
viving in  the  tropics.  Acamptochetus  Cossmann, 
1901  (type  species;  M.  mitraeformis  (Brocchi, 
1814),  Antemetula  Rehder,  1943  (type  species; 
M.  metula  Hinds,  1844)  and  Colubrarina 
Kuroda  and  Habe,  1971  (type  species:  M. 
metulina  Kuroda  and  Habe,  1971)  are  junior 
synonyms. 

Acknowledgments 

I  am  very  much  indebted  to  Ms.  K.  M.  Way  of 
the  Department  of  Zoology,  British  Museum 
(Natural  History)  for  the  loan  of  the  holotype  of 
Buccinum  clathratum.  Dr.  Richard  S.  Houbrick 
of  the  National  Museum  of  Natural  History, 
Smithsonian  Institution  kindly  read  a  draft  of 
the  manuscript  and  allowed  me  access  to  the 
United  States  National  Museum  malacological 
collections.  I  am  indebted  to  Dr.  Richard  H. 
Kilburn  of  the  Natal  Museum  for  reading  the 
manuscript  and  lending  me  a  specimen  critical 
to  this  study.  I  thank  Dr.  Philippe  Bouchet  of  the 
Museum  National  d'Histoire  Naturelle,  Paris  for 
his  comments  on  the  "Metula  problem". 

My  AMNH  colleagues  kindly  provided  valu- 
able contributions  as  follows;  Mr.  Walter  E. 
Sage,  HI,  technical  assistance,  Mr.  Peter  J. 
Harries,  photography,  and  Mrs.  Stephanie 
Grooms,  word  processing. 

I  also  wish  to  join  others  in  congratulating  The 
Nautilus  on  its  100th  anniversary  of  its  found- 
ing and  to  express  my  particular  appreciation 
of  having  known  and  benefited  from  the  friend- 
ship of  three  past  Consulting  Editors- Karl 
Jacobson,  Joe  Rosewater  and  Bill  Clench. 

LITERATURE  CITED 

Adams.  H.  and  A.  18.5.3  [-1854].  The  Genera  of  Recent 
Mollusca,  Arranged  According  to  Their  Organization 
London  l(1.3):65-96,  pis.  9-12  (June,  1853). 

Adams,  A.  and  L.  Reeve.  1850.  Mollusca,  pt.  2:25-44, 
pis.  10-17.  In.  The  Zoology  of  the  Voyage  of  H.M.S. 
"Samarang".  Under  the  Command  of  Captain  Sir  Edward 
Belcher  During  the  Years  1843-1846.  London,  1848-1850. 

Altena,  C.  0.  van  Regteren.  1949.  The  genus.  .Antemetula 
Rehder  in  the  Indo-West  Pacific  area,  with  a  description 
of  two  new  fossil  species.  Bijdragen  Tot  Dierkunde  28: 
385-393.  7  figs. 

Anton.  H.  E.  1839.  Verzeichniss  der  Conchylien  welche  sich 
in  der  Sammlung  von  Hermann  Eduard  Anton  befinden. 
Halle,  xvi  -i-  110  pp. 

Cernohorsky,  W.  O.  1971.  Indo- Pacific  Pisaniinae  (Mollusca: 
Gastropoda)  and  related  buccinid  genera.  Rec.  Auckland 


30      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


Inst,  and  Mus.  8:131-167,  95  figs. 
Cossmann,  M.   1901.  Essais  de  Paleochologie  Comparee, 

pt.  4,  293  pp.,  10  pis. 
Grecchi,  G.  1978.  Problems  connected  with  the  recorded 

occurrence  of  some  mollusks  of  Indo-Pacific  affinity  in 

the  PHocene  of  the  Mediterranean  area.  Riv.  Ital.  Paleont. 

84(3):797-812,  pi.  82,  8  figs. 
Hinds.  R.  B.  1844  [-1845],  Mollusca,  pt.  2,  In  The  Zoology 

of  the  Voyage  of  H.M.S.  Sulphur,  under  the  command  of 

Capt.  Sir  Edward  Belcher  .  .  .  during  1836-1842.  London, 

pp.  25-48.  pis.  8-14  (Oct.  1844). 
Houbrick,  R.  S.  1984.  A  new  "Metula"  species  from  the 

Indo-West    Pacific    (Prosobranchia:    Buccinidae).    Proc. 

Biol.  Soc,  Wash.  97(2):420-424,  1  fig. 
Keen,  A.  M.  1971.  Sea  Shells  of  Tropical  West  America. 

Second  Ed.,  Stanford  Univ.  Press,  viii  -^  1064  pp.,  illus. 
Kiener,    L.   C.    1834   [-1841].   Genre   Buccin.   In.    Species 

general  et  inconographie  des  coquilles  vivantes  .  .  .  Paris, 

vol.  9,  112  pp.,  31  pis. 
Kilburn,  R.  N.  1975.  Taxonomic  notes  on  South  African 

marine  Mollusca  (5):  including  descriptions  of  new  taxa  of 

Rissoidae,  Cerithiidae,  Tonnidae,  Cassididae,  Buccinidae, 

Fasciolariidae,    Turbinellidae,    Turridae,    Architectoni- 

cidae,  Epitoniidae,  Limidae,  Thraciidae.  Ann.  Natal  Mus. 

22(2):577-622,  25  figs. 
Knudsen,  J.  1956.  Marine  prosobranchs  of  tropical  West 

Africa    (Stenoglossa).    Atlantide    Report    No.    4:7-110, 

pis.  1-4. 
Kobelt,    W.    1876    [-1881].    Illustriertes    Conchylienbuch, 

Nurnberg,  vol.  1,  pp.  1-143,  50  plates. 


Kuroda,  T.,  T.  Habe  and  K.  Oyama.  1971.  The  Seashells  of 

Sagami  Bay.   Tokyo,    1-485    +    55  pp.   (English  Text), 

pi.  1-22. 
Malatesta,  A.  1974.  Malacofauna  Pliocenia  Umbra.  Mem. 

Carta  Geol.  Italia.  Rome.  vol.  13.  499  pp.,  32  pis. 
Olsson,  A.  A.  and  F.  M.  Bayer.  1972.  American  metulas 

(Gastropoda:  Buccinidae).  Bull  Mar.  Sci.  22(4):900-925, 

14  figs. 
Rehder,  H.  A.  1943.  New  marine  mollusks  from  the  Antil- 

lean  region.   Proc.   U.   S.   Nat.   Mus.  93(3161):187-203, 

pis.  19,  20. 
Smith,  E.  A.  1904.  On  Mollusca  from  the  Bay  of  Bengal  and 

the  Arabian  Sea.  In.  Series  III.  no.  1.  Natural  History 

Notes  from  H.  M.  Indian  Survey  Steamer  "Investigator", 

Commander  T.  H.  Hemming,  R.  N.  Ann.  Mag.  Nat.  Hist. 

7(13):453-473. 
Tomlin,  J.  R.  le  B.  1927.  The  Mollusca  of  the  "St.  George" 

Expedition.  (I)  The  Pacific  coast  of  S.  America.  Jour. 

Conchology  18(6):  153- 170. 
Vanatta,  E.  G.  1913.  Descriptions  of  new  species  of  marine 

shells.   Proc.  Acad.  Nat.   Sci.   Philadelphia  65(l):22-27, 

3  text  figs.,  pi.  2. 
Woodring,   W.   P.   1928.   Marine  mollusks  from  Bowden, 

Jamaica.   Pt.   2.   Gastropods  and  discussion  of  results. 

Carnegie  Inst.   Washington,   Pub.   no.   385,  vii    +    564, 

pis.  1-4,  3  figs. 
1964.  Geology  and  Paleontology  of  Canal  Zone 

and  adjoining  parts  of  Panama.  Description  of  Tertiary 

mollusks  (Gastropods:  Columbellidae  to  Volutidae).  U.S. 

Geol.  Surv.  Prof.  Pap.  306-C:241-297,  pis.  39-47. 


SOME  ADDITIONAL  TAXONOMIC  UNITS  THAT  FIRST  APPEAR 
IN  PUBLICATIONS  BY  J.  G.  COOPER 

Eugene  Coan 

Research  Associate 

Department  of  Invertebrate  Zoology 

California  Academy  of  Sciences 

San  Francisco,  CA  94118 

ABSTRACT 
Seventeen  taxonomic  units   that  first  appeared  in  publications  by  James 
Graham  Cooper  and  that  were  not  given  in  my  earlier  table  of  his  taxa  are  listed 
here.  All  but  three  are  nomina  nuda. 


It  is  said  that  the  more  complete  a  list  is,  the 
more  glaring  are  its  omissions.  So  it  is,  perhaps, 
with  my  listing  of  the  taxonomic  units  that  first 
appeared  in  the  publications  of  James  Graham 
Cooper  (Coan,  1982:148-185).  Seventeen  names 
that  should  have  been  in  that  list  have  come  to 
my  attention  since  my  book  was  published.  Most 


of  them  were  well  hidden,  being  credited  to 
other  workers.  Fourteen  of  these  names  are 
nomina  nuda,  only  three  actually  being 
validated  by  Cooper.  Two  replace  previous 
homonyms.  The  third  is  a  synonymous  generic 
unit.  It  is  not  impossible  that  other  similarly 
hidden  taxa  will  come  to  light  in  the  future. 


Vol.  lOU(l) 


January  31,  1986 


THE  NAUTILUS      31 


In  the  listing  of  mollusks  below,  I  follow  the 
format  used  in  my  previous  table.  The  Cooper 
papers  cited  refer  to  the  Literature  Cited  of  this 
article,  but  I  also  provide,  in  brackets, 
references  keyed  to  the  complete  bibliography 
of  Cooper's  papers  given  in  my  book  (Coan, 
1982:223-281). 

Class  Bivalvia 

heringi.  Thracia  -  Cooper,  1894  [18;)4b|:  Suppl.,  12th  sheet, 
ex  Dall  MS  [nomen  nuduvi].  Later  validated  by  Dall,  1915: 
442. 
compacta.  Sanguinolaria  -  Cooper,  1894  [1894b]:   126th 

sheet  [nowen  nudum].  Not  subsequently  validated. 
romijlectens.  Tellina  -  Cooper,  1894  [1894b]:  Add.  &  Corr., 
4th  sheet,  ex  Carpenter  MS  [nomev  nudum].  Not  sub- 
sequently validated. 
Philohrya  -  Cooper,  1867  [1867b]:  12,  ex  Carpenter  MS. 
Type  Species  -  Bryophila  .-ietosii  Carpenter,  1864a:  314, 
by  monotypy. 

Remarks  -  Bi-yophila  Carpenter,  1864a:314,  proved  to  be 
a  homonym  (reon  Treitschke,  1825;.57),  and  Carpenter  him- 
self later  renamed  it  as  P/ii7obn/a  (Carpenter,  1872:  index,  p. 
21).  By  then,  however,  it  was  too  late;  Cooper  had  already 
validated  the  name.  Cooper  did  not  expressly  propose 
Philohrya  as  a  replacement  name  and  it  must  therefore  be 
taken  as  a  new  genus. 

Class  Gastropoda 

bimaculata  Clypidella  -  Cooper,  1867  11867b]:  24,  ex  Dall 
MS  ]noinen  nudum].  Later  validated  by  Dall,  1871:132; 
160;  pit.  15,  fig.  7),  as  Fissurellidaea  bimaculata. 

callomarginata.  Clypidella  -  Cooper,  1867  [1867b]:  24,  ex 
Carpenter  MS  ]nomen  nudum].  Later  validated  by  Dall, 
1871:133;  160;  pit.  15,  fig.  8. 

clathftita.  Ocinebra  interfossa  "var."  -  Cooper,  1870 
[1870f]:69,  ex  Carpenter  MS  [nomen  nudum].  Later  vali- 
dated by  Dall,  1919:334,  as  Tritonalia  interjmsa 
clathrata. 

compactum.  Buccinum  -  Cooper,  1894  ]1894b]:  15th  sheet, 
ex  Dall  MS  [noinen  nudum].  Not  subsequently  validated. 

gabbiana.  Chemnitzia  -  Cooper,  1867  [1867b]:  34.  New 
name  for  Turbonilla  graciUiyna  Gabb,  1865:186,  non 
Chemnitzia  gracillima  Carpenter,  1857:431. 

hai'pa.  Tomatina  -  Cooper,  1870  [1870tl:  56,  ex  Dall  MS 
]nomen  nudum].  Later  validated  by  Dall,  1871:136-137; 
160:  pit.  15,  fig.  11. 

Leptothyra  Cooper,  1867  [1867b]:  25.  Type  Species  -  "L. 
sanguinea  Cpr.,"   =    Turbo  sanguinewi  Linnaeus,   1758: 
763  (subsequent  designation  herein). 
Remarks    -    A    synonym   of  Homalapoma   Carpenter, 

1864b:537,  588,  627,  652,  which  has  the  same  type  species. 

Previous  workers  have  evidently  missed  Cooper's  validation 

of  Leptothyra.  dating  it  instead  from  Dall,  1871:130.  It  thus 

becomes  a  senior  homonym  of  Leptothyra  Pease,  1869:70, 

and  if  workers  wish  to  save  this  name,  a  petiton  would  have 

to  be  filed  with  the  International  Commission  on  Zoological 

Nomenclature. 

muricata,    Ocinehra    interfossa   "var."    -    Cooper,    1870 


[1870f]:  69,  ex  Carpenter  MS  ]nomi-u  riuduin].  Not  sub- 
sequently validated. 

paueieostata.  Leptothyra  -  Cooper,  1867  [1867b]:  25,  ex 
Dall  MS  [nomen  nudum].  Later  validated  by  Dall,  1871: 
131;  160;  pit.  15,  fig.  10. 

sitkensis,  Odostomia  -  Cooper,  1894  [1894b]:  100th  sheet, 
ex  Dall  MS  [nomen  nudum].  Not  subsequently  validated. 

steamsiana.  La.mel.la.rin  -  Cooper,  1870  [1870f]:  67,  ex 
Dall  MS  [nomen  nudum].  Later  validated  by  Dall,  1871: 
122;  160;  pit.  15,  fig.  6,  but  as  L.  utearnsii. 

Class  Polyplacophora 

fimbriatus,   Callochilun    -    Cooper,    1867  [lS67b]:  23,  ex. 

Carpenter  MS  [ncmwn  nudum].  Later  validated  by  Keep, 

1887:112,  as  Callistochiton  fimbriatus. 
latior.  Planphorella  -  Cooper,  1894  [1894b]:  113th  sheet, 

ex   Carpenter   MS    [nomen    nudum].    Not   subsequently 

validated. 

LITERATURE  CITED 

Carpenter,  Philip  Pearsall.  1857.  Catalogue  of  the  collection 
of  Mazatlan  shells,  in  the  British  Museum:  collected  by 
Frederick  Reigen  .  .  .  London  (British  Museum)  i-iv  +  ix- 
xvi  4-  552  pp.  (1  Aug.  1857)  [Warrington  ed.,  publ. 
simultaneously]  [reprinted,  Paleo.  Resh.  Inst.,  1967]. 

1864a.  Diagnoses  of  new  forms  of  mollusks  col- 
lected at  Cape  St.  Lucas  by  Mr.  Xantus.  Ann.  Mag.  Natur. 
Hist.  (3)  13(76):31 1-315  (April);  (78):474-479  (June); 
14(79):4.5-49  (.July)  [reprinted  in  Carpenter,  1872: 
207-221]. 

1864b.  Supplementary  report  on  the  present 


state  of  our  knowledge  with  regard  to  the  Mollusca  of  the 
west  coast  of  North  America.  Brit.  Assn.  .Adv.  Sci..  Rept. 
33[for  1863]:517-686  (post-1   Aug.    1864)  [reprinted  in 
Carpenter,  1872:1-172]. 
1872.  The  mollusks  of  western  North  America. 


Embracing  the  second  report  made  to  the  British  Asso- 
ciation on  this  subject,  with  other  papers:  reprinted  by 
permission,  with  a  general  index.  Smithsonian  Inst. 
Misc.  Colin.  10(252):xii  +  325  +  13-121  pp.  (Dec.  1872). 

Coan,  Eugene  V.  1982.  James  Graham  Cooper;  pioneer 
western  naturalist.  Moscow,  Idaho  (Univ.  Press  of  Idaho) 
255  pp.;  31  figs.  (22  Feb.  1982). 

Cooper,  James  Graham.  1867  [1867b  in  Coan,  1982].  Geo- 
graphical catalogue  of  the  Mollusca  found  west  of  the 
Rocky  Mountains,  between  latitudes  33°  and  49°  north. 
San  Francisco,  Calif.  (Geol.  Survey  of  Calif.)  40  pp.  (post- 
April  1867). 

1870  [1870f  in  Coan,  1982].  Notes  on  the  Mol- 
lusca of  Monterey,  California.  Amer.  Journ.  Conch.  6(1): 
42-70  (7  July  1870). 

1894  [1894b  in  Coan,  1982].  Catalogue  of  west 


North  American  and  many  foreign  shells,  with  their  geo- 
graphical ranges.  For  labels,  exchange,  and  check  lists. 
With  a  supplement.  Sacramento,  Calif.  (Calif.  State 
Mining  Bureau)  181  unnumbered  pages  grouped  in  stapled 
sections  [see  Coan,  1982,  for  details],  main  body:  Title 
page  +  Preface  -i-  16(1  pp.;  Siippl.:  Title  page  +  13  pp.; 
Add.  &  Corr.:  5  pp.  (April  1894). 
Dali,  William  Healey.  1871.  Descriptions  of  sixty  new  forms 
of  mollusks  from  the  west  coast  of  North  America  and 


32      THE  NAUTILllS 


January  31,  1986 


Vol.  ]()0(1) 


the  North  Pacific  Ocean,  with  notes  on  others  already 
described.  Amer.  Journ.  Conch.  7(2):9,3-160;  pits.  13-16 
(2  Nov.  1871). 
1915.  A  review  of  some  bivalve  shells  of  the 


group  Anatinacea  from  the  west  coast  of  America.  United 
States  Nation.  Mus.,  Proc.  49(21 16):441-4.56  (27  Nov. 
1915). 

1919.  Descriptions  of  new  species  of  Mollusca 


from  the  North  Pacific  Ocean  in  the  collection  of  the 
United  States  National  Museum.  United  States  Nation. 
Mus.,  Proc.  56(2295):293-371  (30  Aug.  1919). 
Gabb,  William  More.  1865.  Descriptions  of  new  species  of 
marine  shells  from  the  coast  of  California.  Calif.  Acad. 
Sci.,  Proc.  (1)  3(3):182-190  (.Ian.  186.'3). 


Keep,  .Josiah.  1887.  West  coast  shells.  A  familiar  descrip- 
tion of  the  marine,  fresh  water,  and  land  mollusks  of  the 
United  States,  found  west  of  the  Rocky  Mountains.  San 
Francisco  (Bancroft  Bros)  230  pp.;  182  figs.;  frontis. 
(post-July  1887). 

Linnaeus,  Carolus.  1758.  Systema  naturae  per  regna  tria 
naturae  .  .  .  editio  decima,  reformata  1  [Regnum  animale], 
Stockholm  (Laurentii  Salvii).  824  +  iii  pp. 

Pease,  William  Harper.  1869.  Descriptions  of  new  species  of 
marine  gasteropodae  inhabiting  Polynesia.  Amer.  J. 
Conch.  5(2):64-79  (7  Oct.  1869). 

Treitschke,  Friedrich.  1825-1826.  Die  Schmetterlinge  von 
Europa  5.  Leipzig  (G.  Fleischer)  [pts.  1,  2  -  1825;  pt.  3  - 
1826). 


NOTES  ON  THE  HABITS  AND  ANATOMY  OF  THE  INTRODUCED  LAND 
SNAILS,  RUMINA  AND  LAMELLAXIS  (SUBULINIDAE) 

Dee  S.  Dundee^ 

Department  of  Biological  Sciences 

University  of  New  Orleans 

New  Orleans,  LA  70148 


ABSTRACT 

Marking  experiments  showed  that  Rumina  decollata  moved  no  more  than  25 
inches  from  points  of  release  in  a  six-month  period,  demon.'it  rating,  as  with  most 
introduced  forms,  that  passive  distribution  prevails.  Egg-to-egg  span  was  slightly 
over  a  year  in  the  lab.  While  normally  considered  carnivorous,  R.  decollata  ivill 
feed  on  a  variety  of  vegetation,  and  were  routinely  fed  lettuce.  Average  growth  is 
l.J,6  mm,  in  width/ week  and  average  whorl  addition  is  1.18  whorls/week.  At  10  mm 
high,  the  snails  lose  the  upper  3. 0-3.. 5  whorls  and  fill  in  the  top  with  a  .'<eptum.  An 
individual  snail  may  deposit  30-50  ova  over  a  period,  of  several  days  on  more  than 
one  occasion  from  February  to  June  in  this  area.  They  are  deposited  in  shallow 
depressions,  and  soil  particles  adhere  to  them,  thus  rendering  them  in- 
conspicuous. 

Notes  on  the  anatomy  and  a  brief  histological  statement  of  the  digestive, 
nervous,  and  reproductive  systems  o/Lamellaxis  gracilis  are  given. 


Since  the  publication  (Dundee,  1970)  of  a 
study  of  four  introduced  mollusks,  further  infor- 
mation has  been  gathered  about  the  two  species: 
Rumina  decollata  (Linnaeus,  1758)  and  Laniel- 
laxis  gracilis  (Hutton,  1834).  The  information  is 
presented  here  as  follows:  (a)  observations  on 
Rumina  decollata:  movement,  reproduction, 
growth  and  decollation,  longevity  (b)  notes  on 
the  anatomy-histology  of  Lamellaxis  gracilis. 


'Published  posthumously.   Dr. 
Article  proofed  by  the  editor. 


Dundee  died  May  7,   1985. 


Observations  on  Rumina  decollata 
(Linnaeus,  1758) 
Movements.  Six  marking  experiments,  iden- 
tical to  those  described  for  Bradybaena  simi- 
lar is  and  Lamellaxis  gracilis  (see  Dundee, 
1970),  were  done  in  Greenwood  Cemetery  in 
New  Orleans.  The  maximum  movement  demon- 
strated in  any  of  the  experiments  was  25  inches 
from  the  point  of  release  over  a  six  month 
period.  This  seems  to  verify  the  idea  that  in- 
troduced mollusks  do  not  disperse  through  their 
own  power  but,  rather,  are  distributed  through 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS      33 


passive  means. 

Reproduction.  Despite  Sealander's  (1974) 
statement  concerning  self-fertilization  being  ". . . 
the  major  mode  of  reproduction  .  .  ."  in  Rum  inn 
decollata.  Batts  (1957)  reported,  and  I  have 
observed,  regular  matings  between  individual 
snails  thus  suggesting  that  cross-fertilization  is 
the  prevailing  method  in  the  species.  Sealander 
(1974),  later  in  the  paper,  states  that  ".  .  .  we 
cannot  unquivocally  conclude  that  reproduction 
in  the  singles  involved  self-fertilization  rather 
than  parthenogenesis." 

Ova  were  seen  in  both  in  the  field  and  in  the 
lab.  Rascop  (1960)  reported  one  snail  producing 
55-1-  ova  over  a  period  of  3-5  days.  In  my  lab 
colony,  the  members  of  which  began  reproduc- 
ing after  reaching  about  ten  months  of  age,  the 
average  number  of  eggs/snail/deposition  period 
was  32.  In  the  field,  clusters  of  15-41  eggs  were 
observed  at  various  times.  Sealander  (1974),  on 
the  other  hand,  reported  105  snails  laying  6822 
eggs  or,  when  calculated,  64/eggs/snail  average 
which  is  considerably  higher  than  these  aver- 
ages. It  appears  that,  when  one  considers 
Rascop  (1960),  Batts  (1957),  Sealander  (1974), 
and  this  present  work,  that  Rumina  decollata 
may  produce  anywhere  from  7-64  eggs  at  one 
deposition  period  and  then  repeat  the  process 
several  times  over  a  period  of  a  few  weeks. 

The  ova  are  deposited  singly,  but  close  enough 
together  so  that  they  form  clusters.  They  are 
placed  in  shallow  depressions  in  the  soil.  Deposi- 
tion occurs  from  February  to  June  in  this  Gulf 
Coast  area.  A  non-productive  period  then  oc- 
curs, followed  by  more  egg  deposition  from 
September  to  November. 

Average  incubation  time  was  ten  days  at  an 
ambient  temperature  range  of  21-27°C  in  the 
lab.  Batts  (1957)  reported  an  average  time  of 
28.5  days  but  recorded  no  temperatures.  She 
also  reported  one  incubation  time  of  9  days. 
Sealander  (1974)  reported  the  incubation  period 
under  laboratory  conditions  to  be  about  30  days. 
He  did  not  describe  lab  conditions. 

Growth  and  decollation.  Nineteen  lab  juve- 
niles were  observed  for  slightly  over  a  year.  The 
snails,  being  herbivores  as  well  as  predatory  car- 
nivores, were  given  lettuce  and  other  local  snails 
as  food.  The  graph  (Fig.  1)  in  showing  constant 
growth,  demonstrates  indirectly  that  this  is  an 
adequate  diet  to  support  them  since  they  grow 


to  maximum  size  on  it. 

Average  growth  in  height  in  the  lab  at  21- 
27°C  is  1.46  mm/week  and  average  whorl  addi- 
tion is  1.18  whorls/week.  Those  figures  remain 
valid  until  the  snails  reach  10  mm  in  length.  At 
that  10-mm  point  the  upper  3.0-3.5  whorls  are 
lost  and  the  snail  is  said  to  be  "decollate".  If  one 


I 
o 


i 

1 .. 

26|    - 
25 

1 

■ 

—  T  : : : 

'   ■  -   ■   ■ 

l:\'l\^. 

* 

-  •— 

2A 
23 

* 

1 
22 

.  .  .  .  ;     .  • 

'!!!"♦ 

21 
20 

■    ■    i    ■ 

« 
1 

1 

..  .t. ....... .  , , . 

iy 

^ .  1 :  ■ :  ; 

18 

« 

1 

17 

1  .  .  , 

1  .  .  .  . 

* 

..;.... 

.  .  ,.    . 

•     . . 

16- 

,,.... 

1     ■        1        ■  •  1     ■  ■     1 

i  -.!•    ;  •:  1  ■ 

15   ; 

14 

•     ■     1     ■ 

: . : :  i . 
....  1 .    , 

t  ■ 

. 

13 

— -f 

^_^^ 

— 

1 

12  ^  ■ 

11    ^ 

.    1                 t 

-  . 

10    ; 

.         ]     .     .     .     . 

' ::  .1 . 

« 

. 

>    9 

_  ^    .     i 

ue. 

9 

-      -     t     •     -     4      ■ 

-  .  .  (       .... 

^i:;j^;iJ 

1 

8 

1      '      ■ 

' .  flit 

7-  ;j- 

-':'-'[  ; 

K  -  - 

'.^ 

^:: 

-i 

»  t4 

■  ■     -1 

^ 

« 
« 

'                   ' 

-jt?^ 

-  -.<»E 

!  '  1 

+     4.     ^   *- 

TI  I--. 

^ 

.:^..:.|Ti 

41 

^ 

""' 

1 

J: 

: 

1 

^ 

_, 

9    . 

^- 

^- 



_j.^j_4- 

--.-,  , 

— 1 

n- 

-+- 

-1— J— 

-AU- 

i|ilij^ 

'  ''^' 

^ 

^ 

F 

^ 

UI^ 

^ 

WHO  R        L  S 

FIG.  1.  Rumina  ikcollnta.  This  graph  shows  in  terms  of 
height  and  number  of  whorls.  At  the  10  mm  high  point,  the 
snails  become  decollated  thus  having  fewer  whorls  but  con- 
tinuing growth  in  height.  Open  circles  represent  non- 
decollate  individuals;  solid  dots  represent  decollate  ones. 


34      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


examines  a  9-mm-long,  non-decollate  specimen 
(Fig.  2),  one  notices  a  peculiar-appearing  apex 
area.  The  first  3.0-3.5  whorls  have  a  smooth, 
glassy  appearance,  whereas  the  remaining 
whorls  have  some  striations.  The  second  major 
whorl  below  the  protoconch  is  inflated  thus  giv- 
ing the  top  of  the  shell  a  ))uffy  appearance.  The 
shell  decollates  just  l)eneath  the  whorl  below  the 
inflated  one.  I  have  been  unable  to  observe  how 
the  breakage  occurs  although  it  has  been  re- 
ported that  the  snail  bangs  its  upper  whorls 
violently  against  a  hard  object  to  get  rid  of  them 
(Cooke,  189,5).  Batts  (1957)  reported  observing 
them  striking  the  shell  against  rocks  or  the 
aquarium  wall  to  aid  in  decollation.  A  calcareous 
septum,  resembling  the  protoconch,  is  formed 
as  a  cover  for  the  top  of  the  broken  shell  (Fig.  3). 
It  is  easily  seen  in  the  decollate  forms.  If  one 
breaks  off  the  top  whorls  of  a  9.5-mm-long  speci- 
men, one  discovers  that  the  top  3.0-3.5  whorls 
break  much  easier  than  those  below  and  that 
there  is  no  septum  present;  it  is  apparently 
secreted  after  decollation  occurs.  Figure  3  is  a 
view  of  a  septum  as  one  looks  down  on  the  top  of 
a  decollate  shell.  Notice  that  the  umbilicus  is 
retained. 

Decollation  occurs  in  widely  separated  taxa 
from  pulmonates  to  prosobranchs.  It  occurs  in 
terrestrial  as  well  as  aquatic  forms  (brackish 
water).  It  seems  to  have  evolved  independently 
several  times.  Examination  of  a  serial  section  of 
a  9-mm-long  R.  decollata  reveals  nothing 
unusual  about  the  internal  anatomy  in  the  area 
which  is  destined  to  break  off.  One  can  only 
speculate  at  this  point  on  the  selective  advan- 
tage of  casting  off  part  of  the  shell.  Perhaps  the 
upper  whorls  serve  as  a  storage  for  toxins  dur- 
ing early  growth;  perhaps  decollation  is  a 
behavorial  matter  related  to  habitat  (e.g.,  ability 
to  get  under  shelter);  perhaps  it  is  only  a  non- 
selective mutant.  The  entire  matter  needs  fur- 
ther study. 

Longevity.  The  longest  I  have  l)een  able  to 
keep  this  species  alive  in  the  lab  is  just  over  a 
year.  Rascop  (1960)  reports  a  longevity  record 
of  12  years  but  does  not  cite  a  reference.  That 
seems  a  very  long  time  considering  longevities 
of  various  other  snails  (Hyman,  1967).  Years  of 
observations  by  the  author  have  shown  that 
mollusks  tend  to  reproduce  just  prior  to  the  ex- 
tinction of  the  colony.  This  colony,  which  was 


followed  from  egg-to-egg,  began  reproducing  at 
ten  months  which  was  approximately  two 
months  prior  to  the  death  of  the  last  colony 
member. 

Observations  on  Lamellaxis  gracilis 
Shell.  Figure  4  shows  a  typical  adult  shell. 
The  maximum  size  seen  in  the  New  Orleans  area 


F'UiS.  Z  and  3.  Riiniiiid  di'mUata.  2,  A  spi'cmifn  of  ;i|j|jnix- 
irriatoly  9  mm  showing  the  smooth  quahly  of  the  top  three 
whorls  as  opposed  to  the  striations  on  the  remaining  ones.  3, 
A  view  into  the  top  of  a  large  decollate  specimen. 


Vol.  100(1) 


January  31,  1986 


THE  NAUTILUS 


FIGS.  4-7.  Lamelhxif:  gracili.'i.  4,  Adult  shell. 
5,  An  about-to-hatch  snail  dissected  from  its  egg 
case.  6,  The  jaw,  7,  Central  and  lateral  teeth. 

was  13  mm  in  length,  3.5  mm  in  width  at  the 
broadest  point,  and  with  9.25  whorls.  The  maxi- 
mum size  of  Lamellaxis  micra  (Orbigny,  1835) 


which  also  has  been  introduced  to  this  area  was, 
by  contrast,  7.2  mm  x  2.2  mm,  and  had  6.75 
whorls.  The  color  of  L.  gracilis  is  light  tan  and 
all  whorls  except  the  protoconch  are  finely 
striated.  A  small  umltilicus  is  partially  hidden  by 
a  minor  reflection  of  the  lip  (Fig.  4).  Unhatched 
ova  (Fig.  5)  removed  from  the  female  system 
show  clearly  the  beginnings  of  striation  except 
on  the  protoconch. 

Jaw.  This  structure  is  found  at  the  entrance 
to  the  pharynx  in  a  mid-dorsal  position.  As  can 
be  seen  in  Figure  6,  it  is  a  single,  curved  device 
which  has  coarse  transverse  ribbing. 

Radula.  The  radula  (Fig.  7)  is  relatively  nar- 
row with  little  variation  among  central,  lateral, 
and  marginal  teeth.  The  teeth  are  reminiscent 
of  marginals  of  Helix  aspersa  except  the  lateral 
cusps  are  more  pronounced. 

Digestive  system.  The  esophagiis,  lined  by 
tall,  ciliated  columnar  epithelium  and  underlain 
by  a  thin  circular  muscle  layer  surrounding  an 
equally  thin  longitudinal  layer.  The  esophagus 
leaves  the  pharyngeal  area,  passes  through  the 
nerve  ring  and  posteriorly  into  the  stomach; 
(Fig.  9  ST).  In  keeping  with  other  Stylommato- 
phora,  no  crop  was  found.  The  stomach  is  lined 
by  tall,  columnar,  ciliated  epithelium  underlain 
by  a  double  muscle  layer  (longitudinal  and  cir- 
cular) and  surrounded  by  connective  tissue.  The 
lining  is  folded  in  several  areas.  The  digestive 
gland  (Fig.  8,  D)  empties  into  the  stomach  via 
two  ducts -one  from  each  section  of  gland.  A 
dissection  showing  the  two  parts  is  very  difficult 
and  can  only  be  done  with  fresh  (non-preserved) 
material.  Histological  observations  reveal  two 
cell  types  making  up  the  digestive  gland  (also 
called  liver  and  hepatopancreas).  A  tall  colum- 
nar cell  type  which  contains  numerous  vacuoles 
is  often  seen  with  the  lumen  in  a  state  of  disinte- 
gration implying  secretion.  The  second  type  is  a 
low,  non-ciliated  triangular  form.  Carriker  and 
Bilstad  (1946)  reported  similar  cells  and  called 
them  "digestive"  and  "calciferous"  and  deter- 
mined that  the  latter  contain  calcium  phosphate 
granules.  Digestion  is  apparently  both  intra- 
and  extra-cellular.  Sections  reveal  partially 
digested  material  in  the  stomach  lumen  (Fig.  8, 
ST)  and  also  in  some  the  cells  of  the  digestive 
gland.  An  intestine  leaves  the  stomach,  loops 
around  in  the  same  whorl  as  the  stomach, 
descends  to  the  right  dorso-lateral  of  the  head- 


36      THE  NAUTILUS 


January  31,  1986 


Vol.  100(1) 


FIG.  8.  Lamt'lliiji.^  ^innms.  Parasagittal  section  of  a 
young  snail.  D  =  digestive  gland;  ST=  stomach;  I  =  intestine; 
A  =  albumen  gland. 


foot  where  the  anus  is  located.  The  intestine 
(Fig.  8,  I)  is  lined  by  tall,  ciliated  columnar 
epithelium  containing  mucous  cells  and  under- 
lain by  an  inner  longitudinal  and  an  outer  cir- 
cular muscle  layer. 

Nervous  system.  The  ganglia  form  a  ring 
around  the  anterior  portion  of  the  digestive 
tract  just  posterior  to  the  pharynx.  This  is  the 
result  of  detorsion  wherein  the  parietal  (in- 
testinal) and  visceral  ganglia  migrate  forward 
near  the  pleural  ganglia.  In  Lamellaxis  the  end 
result  is  two  cerebral  ganglia  on  the  dorsal  of 
the  ring  and  two  pleurals,  two  parietals,  and  a 
visceral  (two  fused  ganglia)  forming  the  ventral 
portion  of  the  ring  and  innervating  the  entire 
visceral  mass.  The  total  nervous  system  was  not 
traced. 

A  pair  of  eyes,  located  on  the  dorsal  tips  of  the 
posterior  tentacles,  are  typical  pulmonate  vesi- 
cular types  (Hyman,  1967,  p.  584).  The  stato- 
cysts,  located  adjacent  to  the  pedal  ganglia  in 
the  foot,  are  small  spherical  vesicles  each  con- 
taining calcareous  granules.  Each  is  covered  by 
a  layer  of  connective  tissue  and  lined  internally 


by  a  squamous-type  epithelium  with  a  layer  of 
cilia  equally  spaced  around  the  inner  periphery. 

Excretory  system.  A  single  nephridium  is 
located  near  the  dorso-posterior  of  the  pulmon- 
ary cavity.  If  one  holds  the  snail  so  that  the 
aperture  is  facing  him,  the  reno-pericardial 
mass  is  on  the  extreme  lower  right  of  the  whorl 
above  the  aperture.  The  pericardial  cavity  has  a 
duct  from  it  into  the  sac-like  nephridium.  That 
nephridium,  in  most  Pulmonates,  opens  into  the 
|.)ulmonary  cavity  directly  via  a  slit  or  a  short 
ureter.  I  could  not  determine  which  is  present  in 
this  snail  either  by  dissection  or  sections. 

Reproductive. System.  Lamellaxis  is  herma- 
phroditic. The  ovotestis,  a  grapelike  cluster 
embedded  in  the  digestive  gland  proximal  to  the 
columella,  contains  both  oocytes  near  the  peri- 
phery of  each  follicle  and  spermatocytes  and 
sperm  more  centrally  located  near  the  duct  of 
each.  A  coiled  hermaphroditic  duct,  lined  by  low 
ciliated  cuboidal  epithelium,  drains  both  the 
ovary  and  testis.  At  the  lower  end  of  that  duct  a 
small  saclike  structure,  the  seminal  vesicle,  ap- 
pears to  branch  off  and  serves  as  a  sperm  stor- 
age area.  It  is  seen  full  of  sperm  during  the 
breeding  season.  Near  that  point,  the  male  and 
female  systems  part  company  with  each  system 
continuing  downward,  closely  applied  to  and 
paralleling  each  other  (spermoviduct  of  some 
authors)  through  the  whorls  towards  the 
anterior  end. 

The  male  system  consists  of  a  sperm  duct  (vas 
deferens)  with  a  ciliated  cuboidal  lining.  It 
receives  a  prostatic  secretion  via  a  small  connec- 
tion between  the  prostate  and  vas  deferens.  The 
prostate  consists  of  cuboidal  cells  around  an  ir- 
regular lumen.  The  cells  closest  to  the  lumen 
disintegrate  as  a  result  of  secretion.  Nuclei  are 
basal.  The  vas  deferens  continues  downward 
and  enters  the  verge.  Just  prior  to  that  entry 
the  duct  liecomes  lined  by  tall,  ciliated  glandular 
epithelium.  The  system  empties  on  the  right  side 
of  the  head  near  the  base  of  the  right  tentacle. 
The  verge  is  simply  a  long  attenuated  tubule 
with  no  secondary  structures  on  it.  At  the  point 
where  the  systems  become  separate  the  oviduct 
appears  to  enter  a  large  albumen  gland  but, 
actually,  the  gland  is  large  enough  that  it  has 
the  oviduct  "embedded  in"  it  (Fig.  9,  A).  The 
allmmen  gland  has  a  varied  histological  struc- 
ture  ranging   from   cuboidal   to   low,   ciliated, 


Vol.  1(H)(1) 


January  31,  1986 


THE  NAUTILUS      37 


epithelial  cells.  This  gland,  in  section,  always  ap- 
pears striated  and  somewhat  "crumbled".  The 
lumen  ends  of  the  cells  appear  to  disintegrate  as 
secretions  are  formed. 

Just  ventral  to  this  albumen  gland,  at  the 
separation  of  the  male-female  systems  is  an  area 
where  fertilization  occurs  (fertilization  pouch  or 
carrefour  in  other  pulmonates:  e.g.,  (Petrellis 
and  Dundee,  1969).  A  distinct  pouch  could  not 
be  located  in  L.  gracilis.  Only  a  slightly  swollen 
area  in  the  hermaphroditic  duct  exists.  It  is  dif- 
ferent histologically  from  the  initial  part  of  the 
duct  in  that  the  epithelium  is  ciliated  columnar 
in  contrast  to  low,  non-ciliated  cuboidal  in  initial 
portion  of  the  duct.  The  change  in  histological 
structure  indicates  that  the  area  is  the  equiva- 
lent of  the  fertilization  pouch  and  is  doubtlessly 
the  site  of  fertilization. 

Fertilized  eggs  receive  their  albumen  coat  as 
they  pass  by  the  albumen  gland.  As  they  con- 
tinue anteriorly  downward,  the  shell  is  added  by 
the  mucus  gland  (also  called  capsule  gland) 
which  is  a  continuation  of  the  large  glandular 
mass.  The  albumen  and  mucus  glands  can  be  dis- 
tinguished microscopically  by  change  from  whit- 
ish, coarse  appearance  (albumen)  to  a  smooth 
yellowish  one  with  the  entire  mass  convoluted 
(mucus).  Microscopically  the  mucus  gland  dif- 
fers from  the  albumen  gland  described  above  by 
being  a  highly  glandular  mass  lined  by  cuboidal 
cells  which,  in  many  cases,  are  secreting  as 
evidenced  by  the  disintegrated  inner  cell  mass. 
Non-secretory  cells  are  ciliated,  and  cell  boun- 
daries are  indistinct. 

Often  as  many  as  6-8  large  ova  can  be  seen 
lined  up  in  the  oviduct  in  the  body  whorl  and  the 


one  immediately  above  it.  Just  at  the  point 
where  ova  emerge  from  near  the  mucus  gland, 
the  vagina  receives  a  duct  from  the  spermatheca 
(also  called  seminal  receptacle  or  bursa),  a  sac 
which  stores  sperm  from  copulation.  Histologi- 
cally the  spermatheca  is  composed  of  non- 
ciliated,  tall  columnar  epithelium  encapsulated 
in  a  muscle-connective  tissue  layer.  The  vagina 
is  a  muscular  tubule  lined  by  ciliated  epithelium. 
It  ends  at  the  gonopore  which  is  found  in  the 
right  dorso-lateral  area  of  the  head. 

LITERATURE  CITED 

Batts,  Jeanne  H.  1957.  Anatomy  and  life  cycle  of  the  .'^nail 
Rumina  decollata.  Southwest  Nat.  2:74-82. 

Biswas,  S.  K..  R.  Rahman  and  T.  R.  Mitra.  1976.  Observa- 
tions on  the  breeding  habits  of  Opeas  gracile  (Hutton) 
(Gastropoda:  SubuMnidae).  .Jour.  Conch.  29:69-70. 

Carriker,  Melbourne  and  N.  M,  Bilstad.  1946.  Histology  of 
the  ahmentary  system  of  the  snail,  Lymnnen  stagnalis 
(ippressa.  Trans.  Amer.  Micr.  Soc.  64:250-275. 

Cooke,  A.  H.  1895.  In  Cambridge  Natural  History  Vol.  HI 
(Molluscs  and  Brachiopods).  Macmiilan  &  Co.  (Reprint 
edition:  1-ix  +  535;  334  figs.;  1  map.  Wesley,  Ltd., 
England,  1959). 

Dundee,  Dee  S.  1970.  Intro<luced  Gulf  Coast  molluscs. 
Tulane  Studies  in  Zool.  and  Bot.  1G(3):101-115. 

Dundee,  Dee  S.  and  Roy  J.  Baerwald.  1983.  Observations  on 
a  micropredator,  Gulella  bicolor  (Hutton),  (Gastropoda: 
Pulnionata:  Streptaxidae).  The  Nautilus  98(2):63-68. 

Hyman,  Libbie  H.  1967.  The  Inverteljrata:  Vol.  VI  Mollusca 
'l:  i-vii  +  1-792;  249  figs.;  McGraw  Hill  Book  Co. 

Petrellis,  Louisa  S.  and  Dee  S.  Dundee.  1969.  Veronicella 
ameghini  (Gastropoda:  Reproductive,  Digestive,  Nervous 
system).  Trans.  Amer.  Micr.  Soc.  88(4):547-558. 

Rascop.  Ann  M.  1960.  The  biology  of  Rumuin  decollata. 
Unpub.  M.A.  Thesis.  Univ.  Ariz. 

Sealander,  R.  K.,  D.  W.  Kaufman  and  S.  Ralin.  1974.  Self- 
fertilization  in  the  terrestrial  snail  Rumina  decollata. 
Veliijer  16(3):265-270. 


New  Awards 

The  Division  of  Mollusks,  Department  of 
Invertebrate  Zoology,  National  Museum  of 
Natural  History,  Smithsonian  Institution 
announces  the  availability  of  two  fellowships  to 
be  awarded  to  graduate  students  of  systematic 
malacology. 

1.  Rosewater  Fellow  Award  (up  to  $500) 

2.  Smithsonian  -  Conchologists  of  America 

Fellow  Award  (up  to  $1,000) 


These  awards  are  to  help  support  students  for 
short  term  research  visits  to  the  collections  and 
libraries  of  the  Division  of  Mollusks,  National 
Museum  of  Natural  History  and  are  to  be  used 
for  systematic  studies  of  Mollusca.  Funds  can 
help  cover  travel,  subsistence,  and  research 
costs  [xerox,  postage,  etc.].  Interested  students 
should  submit  a  1-page  proposal,  a  budget  with 
indication  of  matching  funding,  if  available,  and 
a  supporting  letter  from  their  faculty  advisors. 
Deadline  for  applications  is  March  1,  1986. 
Awards  will  be  announced  on  April  1,  1986. 


38      THE  NAUTILUS 


January  31,  1986 


Vol.  11)0(1) 


NOTES 

NOTE  ON  CRYPTONATICA  DALL,  1892 
(GASTROPODA:  NATICIDAE) 

Richard  E.  Petit 

806  St.  Charles  Rd. 
North  Myrtle  Beach,  SC  29582 

The  naticid  subgenus  Cryptonatica  was  pro- 
posed by  Dall  (1892:362)  with  no  type  designa- 
tion. Dall's  first  mention  of  Cryptonatica  {p.  362) 
lists  the  species  Natica  clausa  Broderip  & 
Sowerby,  A'^.  russa  Gould  and  A'^.  pusilla  Say  as 
being  included.  A  few  pages  later  (p.  366)  the 
new  species  Nafica  (Cryptonatica)  floridana  is 
described.  As  pages  362-366  were  published  at 
the  same  time,  all  four  of  these  taxa  are  original- 
ly included  species  and  are  available  for  designa- 
tion as  type-species  of  Cryptonatica. 

Cossmann  (1896:238)  in  a  review  of  Dall's 
1892  publication  designated  A'^.  (C.)  floridana 
Dall  as  the  type  of  Cryptonatica,  an  action 
overlooked  by  all  subsequent  authors,  including 
Cossmann  himself.  Dall  (1909:85)  designated  A^. 
claMsa  Broderip  &  Sowerby  as  type,  and  this 
species  has  been  accepted  as  type  (although  with 
differing  citations  for  the  designation)  by  all 
subsequent  authors.  Among  those  citing  A^. 
claitsa  as  type  of  Cryptonatica  are  Cossmann 
(1925:120),  Woodring  (1928:384),  Boss, 
Rosewater  &  Ruhoff  (1968:95),  and  Marincovich 
(1977:409). 

Natica.  clausa.  is  a  cold-water  Recent  species 
and  A^.  floridana  is  from  the  Florida  Oligocene. 
It  is  not  the  purpose  of  this  paper  to  investigate 
or  discuss  the  probable  relationship  of  these 
species  or  the  ramifications  possible  as  a  result 
of  Cossmann's  type-designation,  but  to  call  to 
the  attention  of  naticid  systematists  it's  ex- 
istence so  that  appropriate  action  may  be  taken. 

Cossman  often  made  type  designations  in  his 
reviews  which  appeared  in  various  publications 
from  1888  to  1924.  A  listing  of  these  designa- 
tions is  now  in  preparation  and  will  be  published 
in  the  future. 

LITERATURE  CITED 

Bo.ss,  K.  .1.,  ,1.  Rosewater  and  F.  A.  Ruhoff.  1968.  The 
Zoolo)rical  Taxa  of  William  Healey  Dall.  U.S.N.M.  Bull. 
287:1-427. 

Cossmann,  M.  189().  Revue  de  Paleoconcholofjie.  La  Feuillc 
des  Jeunes  Naturalistes  26(:3I2):2:i0-24(). 

1925.   Naticidae.   Essais  de   Paleoconchologie 


Comparee  13:97-150. 
Dall,  W.  H.  1892.  Contributions  to  the  Tertiary  Fauna  of 

Florida  ...  Wagner  Free  Inst.  Sci.  3(2):201-473,  pis.  13-22. 
1909.  The  Miocene  of  Astoria  and  Coos  Bay, 

Oregon.  U.S.  Geol.  Surv..  Prof.  Paper  59:1-278,  pis.  1-23. 
Marincovich,   L.,  Jr.   1977.  Cenozoic  Naticidae  (Mollusca: 

Gastropoda)  of  the   Northeastern   Pacific.   Bull.    .Amer. 

Paleont.  70(294):16.5-494,  pis.  17-42. 
Woodring,  W.  P.   1928.  Miocene  Mollusks  from  Bowden, 

Jamaica.  Pt.  II.  Gastropods  and  Discussion  of  Results. 

Carnegie  Inst.  Wash.  Publ.  385.  564  pp.,  40  pis. 


NATICA  (GLYPHEPITHEMA)  BAYERL 
NEW  NAME 

Harald  A.  Rehder 

National  Museum  of  Natural  History 
\  Smithsonian  Institution 

Washington,  D.C.  20560 

Richard  E.  Petit  has  called  my  attention  to  the 
fact  that  the  species  in  the  Naticidae  that  I 
described  as  Glyphcpithema  floridana  in  1943 
(Proc.  U.S.  National  Museum:  93(3161):196-197, 
pi.  19,  figs.  19-21)  is  preoccupied  by  Natica 
(Cryptonatica)  Jloridana  Dall,  1892,  described 
from  the  Oligocene  of  the  Tampa  formation 
(Trans.  Wagner  Free  Inst.  Sci.  Philadelphia 
3(2):366,  pi.  17,  fig.  5).  The  taxon  Glyphepit he- 
ma.  Rehder,  1943  (op.  cit.,  p.  196),  originally 
proposed  by  me  as  a  genus,  is  now  quite  general- 
ly cited  as  a  subgenus  of  Natica.  and  therefore 
the  species  name  floridana  Rehder  needs  to  be 
replaced. 

I  propose  to  name  this  species  Natica  (Gly- 
phepitheyna)  bayeri  Rehder  dedicating  it  to  my 
longtime  associate  and  friend,  F.  M.  "Ted" 
Bayer,  who  collected  the  holotype  in  Lake 
Worth,  Florida.  The  species  is  found  from  south- 
eastern Florida  to  Panama  and  northeastern 
Brasil  (Abbott,  American  Seashells,  ed.  2,  p. 
159,  1974). 

DEATHS 
A.  Myra  Keen  -  1905  -  1986 

One  of  the  great  giants  of  American  malacology,  Angeline 
Myra  Keen,  died  on  January  4,  1986,  at  the  age  of  80  in 
California.  Dr.  Keen  was  one  of  the  leading  forces  in  the 
development  of  students  in  malacology  at  Stanford  Univer- 
sity, and  well-known  not  only  for  her  excellent  taxonomic 
work  hut  also  for  her  major  contribution  of  popular  books 
that  aided  legions  of  appreciative  private  shell  collectors. 
She  assisted  and  gave  advice  to  many  amateurs  who  wished 
to  publish  on  mollusks. 


INFORMATION  FOR  SUBSCRIBERS 


The  annual  subscription  rate  for  The  Nautilus 
is  $15.00  for  individuals  (foreign  $18.00)  and 
$20.00  for  institutions  (domestic  or  foreign). 
Subscriptions  may  begin  in  January.  Send  check 
or  money  order  made  out  to  "American  Mala- 
cologists"  to  the  Business  Manager,  P.O.  Box 
2255,  Melbourne,  Florida  32902-2255,  U.S.A. 


Back  issv£s  from  volume  72  to  date  are  ob- 
tainable from  the  Business  Manager.  Volumes  1 
through  71  (if  available)  may  be  obtained  in 
reprint  or  original  form  from  Kraus  Reprint 
Co.,  Route  100,  Millwood,  New  York  10546. 

Advertising  rates  may  be  obtained  from  the 
Business  Manager  or  Editor. 


CONTRIBUTORS 


Manuscripts:  Authors  are  requested  to  follow 
the  recommendations  of  the  Style  Manual  for 
Biological  Journals,  which  may  be  purchased 
from  the  American  Institute  of  Biological  Sci- 
ences, 1401  Wilson  Boulevard,  Arlington,  Va. 
22209.  Manuscripts  should  be  typewritten  and 
doublespaced;  original  and  one  copy  are  re- 
quired, to  facilitate  reviews.  Tables,  numbered 
in  arable,  should  be  on  separate  pages,  with  the 
title  at  the  top.  Legends  to  photographs  should 
be  typed  on  separate  sheets.  Explanatory  terms 
and  symbols  within  a  drawing  should  be  neatly 
printed,  or  they  may  be  pencilled  in  on  a  translu- 


cent overlay,  so  that  the  printer  may  set  them  in 
8  pt.  type.  There  is  a  charge  of  50  cents  per 
word  for  this  extra  service.  All  authors  or  their 
institutions  will  be  charged  50  cents  per  line  of 
tabular  material  and  taxonomic  keys.  The  pub- 
lishers reserve  the  right,  seldom  exercised,  to 
charge  $45  per  printed  page. 

An  abstract  should  accompany  each  paper. 

Reprints  are  available  at  cost  to  authors. 
When  proof  is  returned  to  authors,  information 
about  ordering  reprints  will  be  given.  They 
are  obtained  from  Economy  Printing  Co.,  Inc., 
R.D.  3,  Box  169,  Easton,  Maryland  21601-9430. 


MOLLUSK  VOUCHER  SPECIMENS 


It  is  becoming  increasingly  important  for 
future  research  purposes  that  an  identified  sam- 
pling of  species  mentioned  in  publications  be 
deposited  in  a  permanent,  accessible  museum 
specializing  in  mollusks.  This  is  particularly  true 
of  mollusks  used  in  physiological,  medical, 
parasitological,  ecological,  and  experimental 
projects. 


Several  museums  of  natural  history  have  ex- 
tensive modern  facilities  and  equipment  for  the 
housing  and  curating  of  voucher  specimens. 
Material  should  be  accompanied  by  the  identifi- 
cation, locality  data  and  its  bibliographic 
reference.  There  is  no  charge  for  this  perma- 
nent curating  service,  and  catalog  numbers,  if 
desired,  will  be  sent  to  authors  prior  to  publica- 
tion. 


WANTED  -  OLD  SHELL  BOOKS 

Will  pay  good  prices  for  libraries,  second-  Phone  (1-305-725-2260)  or  write:  R.  Tucker 

hand  books  and  reprints  on  mollusks,  shells  Abbott,  Aynerican  Malacologists,  Inc.,  P.O. 

and    conchology.    Back    numbers    of    The  Box  2255,  Melbourne,  FL  32902.  Free  ap- 

Nautilvs,  vols.  40-71  wanted,  $1.50  each.  praisals. 


americanmalacologists,  inc. 

PUBLISHERS  OF  DISTINCTIVE  BOOKS  ON  MOLLUSKS 

THE  NAUTILUS  (Quarterly) 

MONOGRAPHS  OF  MARINE  MOLLUSCA 

STANDARD  CATALOG  OF  SHELLS 

INDEXES  TO  THE  NAUTILUS 

(Geographical,  vols  1-90;  Scientific  Names,  vols  61-90) 

REGISTER  OF  AMERICAN  MALACOLOGISTS 


APRIL  30,  1986 


THE 


NAUTILUS 


ISSN  0028-1344 


Vol.  100 


No.  2 


A  quarterly 

devoted  to 

malacology  and 

the  interests  of 

conchologists 


hWim  Bioiwical  LsbcralOTj'  \ 
LIBRARY  I 

MAY    7  1986 

I     Woods  Hole,  Mass.     ! 


Founded  1889  by  Henry  A.  Pilsbry.  Continued  by  H.  Burrington  Baker. 
Editor-in-Chief:  R.  Tucker  Abbott 


EDITORIAL  COMMITTEE 
CONSULTING  EDITORS 


Dr.  William  K.  Emerson 
Department  of  Living  Invertebrates 
The  American  Museum  of  Natural  History 
New  York,  NY  10024 

Mr.  Samuel  L.  B.  Fuller 
1053  Mapleton  Avenue 
Suffield,  CT  06078 

Dr.  M.  G.  Harasewych 
363  Crescendo  Way 
Silver  Spring,  MD  20901 

Mr.  Richard  L  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Los  Angeles  County  Museum  of  Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 


Dr.  Arthur  S,  Merrill 
c/o  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Donald  R.  Moore 
Division  of  Marine  Geology 
School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 

Dr.  G.  Alan  Solem 
Department  of  Invertebrates 
Field  Museum  of  Natural  History 
Chicago,  IL  60605 

Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Gilbert  L.  Voss 
Division  of  Biology 

School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 


EDITOR-IN-CHIEF 

Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
Box  2255,  Melbourne,  FL  32902-2255 

Mrs.  Cecelia  W.  Abbott 
Business  and  Subscription  Manager 
P.O.  Box  2255 
Melbourne,  FL  32902-2255 


Second  Class  Postage  paid  at  Melbourne,  Florida 
and  other  post  offices 


The  Nautilus  (USPS  374-980) 
ISSN  0028-1344 

A  quarterly  magazine  devoted  to  malacology. 
CopyriKht  ^'ISSe  by  American  Malacolopsts,  Inc. 

OFFICE  OF  PUBLICATION 

American  Malacologists,  Inc.  (United  Parcel  Address: 

2208  South  Colonial  Drive,  Melbourne,  FL  32901) 

Mail:  Box  2255,  Melbourne,  FL  32902-2255 

POSTMASTER:  Send  address  changes  to  above. 

Subscription  Price:  $15.00  (see  inside  back  cover) 

$17.00  (foreign);  institutions  $20.00 


LIBRARY 

MAY    7  1986 

Woods  Hole,  Mass. 


THE 

NAUTILUS 

Volume  100,  number  2  -  April  30,  1986 

ISSN  0028-1344 


CONTENTS 


Thomas  R.  Waller 

A  New  Genus  and  Species  of  Scallop  (Bivalvia:  Pectinidae)  from  off  Somalia, 

and  the  Definition  of  a  New  Tribe  Decatopectinini 39 

Hans  Bertsch  and  Antonio  Mozqueira  Osuna 

A  New  Species  of  Triton  in  (Nudibranchia)  from  Southern  California  and  Baja  California 46 

Arthur  H.  Clarke 

Unionidae  of  the  Upper  Connecticut  River,  a  Vanishing  Resource 49 

Jose  Gomez,  Mercedes  Vargas  and  Emile  A.  Malek 

Morphological  Anomalies  in  the  Shell  of  P"ield-CiilleetedS('(W(/)/;a/(/n'((  glabratn  (Say,  1818).  .  .  .53 

Eva  Pip 

The  Ecology  of  Freshwater  Gastropods  in  the  Central  Canadian  Region 56 

Jose  Gomez,  Mercedes  Vargas  and  Emile  A.  Malek 

Pseudosuccinea  columella  (Lymnaeidae)  in  the  Dominican  Republic  and 

Transmission  of  Fascioliasis  in  the  Caribbean  Region 66 

David  Nicol 

Some  Aspects  of  the  Evolution  of  the  Rudist  Pelecypods 69 

Mark  E.  Gordon 

A  New  Sitmatogyrus  from  the  Southwestern  Ozarks  with  a  Brief  Review 

of  the  Hydrobiidae  from  the  Interior  Highlands  (Gastropoda:  Prosobranchia) 71 

Anthony  D'Attilio  and  Barbara  W.  Myers 

Favartia  brericida  (Sowerby,  1834)  and  Two  New  Species  of  Favurtia 

from  the  Western  Pacific  (Gastropoda:  Muricidae) 78 

New  Mollusk  Journals 55        Meeting  News 68 


SEASHELL 
TREASURES 

or  THE  CARIBBEAn 

text  and  photographs 
LESLEY  SUTTY 

(under  the  editorship  of  R.  Tucker  Abbott) 

Follow  the  personal  adventures  of  a  marine  biolojjist  as  she  hunts  throu^^hout  the  West  Indies 
for  elusive  and  rare  seashells.  An  active  naturalist  and  archaeologist.  Miss  Suttv  has  spent  years 
in  photoj^raphing  and  observinjj  marine  life  and  studying  the  uses  of  shells  by  the  ancient  Carib 
Indians.  Under  the  editorial  guidance  of  Dr.  Abbott  this  book  serves  as  an  accurate  guide  to  the 
most  unusual  shell  treasures  of  the  Caribbean. 


New  Discoveries 

Not  seen  since  first  discovered  35  years  iigo.  Straus's 
Murex  (Hexaplex  strausi  Verrill,  1950)  was  found  alive 
in  21  feet  of  water  in  Martinique.  Lesley  re-discovered 
Lamarck's  long-lost  Picrynoius plnllopwnis  as  early  as 
1469.  and  she  is  the  first  to  record  a  case  of  a  human 
being  slung  by  a  West  Indian  poisonous  cone  shell. 

Deep  Dive  Photos 

For  the  first  time,  a  living  specimen  ot  the  rare 
Surinam  Cowrie  is  photographed  during  a  night  scuba 
dive.  See  an  eyeball-to-eyeball  photo  ot  the  queer 
Umbrella  Shell  (Umhracuhtm).  and  thrill  to  the  color 
shots  of  intricate  niulihranch  sea  slims. 


A  UNIQUE  SHELL  BOOK! 

This  unusual  book  is  a  blend  of  beauty, 
adventure  and  science: 

/.    11)0  rare  spccu-s  illuslnilVit  in  i^oiin'ous  color. 
2.    Details  of  where  llwv  live  and  how  lo  find  them. 
.?.   SeieiUijie  and  eoinnion  names,  each  nitli  author  and  dale. 
4.    (ieoi;ra/)hieal  oceiirreme.  depth  and  relative  abiindancv. 
5.   Colleclini;  experiences  ihroii^lunii  the  .\nlilles. 
6,    I'aeked  with  /icir  hiolofiieal  ohser\ati(nis. 

May  1986.  cloth 

NATURAL  HISTOKY/COLLCCTinQ      138  color  plates 

Bibliography  Index  7'/2  x  9%"     128  pages 


EXCLUSIVE  OFFER! 

A  handsome  bookplate  has  been  designed 
for  the  maiden  publication  of  SEASHELL 
TREASURES  OF  THE  CARHJBEAN. 
Your  numbered  copy  with  your  name, 
will  be  personally  autographed  by  the 
author.  Miss  Lesley  Sutty  of  the 
Grenadine  Islands. 


RESERVE  YOUR  COPY  NOW! 
Only  $22.95 

SEND  TO: 

American  Malacologists 

(New  Enjjland  Division) 

P.O.  Box  1192,  BruLlNCTON,  MA  ()1,S()3 


Personal i/ed  Bookplate 
Numbered  Book 


Lach    copy    is    assigned    a    luimbei     as    orders    arc 
received.  We  guarantee  dcliverv  or  a  lull  refund 


Vol.  100(2) 


A()ril  30,  1986 


THE  NAUTILUS      39 


A  NEW  GENUS  AND  SPECIES  OF  SCALLOP 

(BIVALVIA:  PECTINIDAE)  FROM  OFF  SOMALIA, 

AND  THE  DEFINITION  OF  A  NEW  TRIBE  DECATOPECTININI 


Thomas  R.  Waller 

Dt'pai'ttiifiit  (if  Paleobiology 

U.  S.  National  Museum  of  Natural  History 

Smithsonian  Institution 

Washing-ton,  DC  205(i0 

ABSTRACT 

Somalipecten  Waller,  new  genus,  is  one  often  extant  pectinid  genera  that  are 
placed  in  the  new  tribe  Decatopectinini,  which  is  characterized  by  low  early  beak 
inflation,  very  closely  spaced  commarginal  lamellae,  and  very  weak  or  absent 
resilial  hinge  teeth.  The  type  species,  Somalipecten  cranmerorum,  new  species, 
from  offSom.alia  in  1.50-300  m.  is  the  sole  survivor  of  a  group  of  congeneric  species 
that  lived  in  the  Neogene  and  Quaternary  if  the  Red  Sea  and  we.^terti  Indian 
Ocean  regions. 


Introduction 

Many  new  species  of  shelled  moilusks  are  still 
being  discovered  in  the  world's  oceans,  but  it  is 
remarkable  when  a  new  species  turns  out  to  be 
large  in  size,  exquisitely  beautiful  in  color  and 
form,  and  also  abundant  at  its  type  locality  in  a 
region  thought  to  have  a  well-known  moUuscan 
fauna.  In  this  report  I  describe  a  new  scallop, 
Somalipecten  cranmerorum.  new  genus  and 
species,  taken  by  Taiwanese  trawlers  working 
new  fishing  grounds  off  Somalia.  This  new  pec- 
tinid genus,  which  has  an  extensive  fossil  record 
in  Neogene  and  Quaternary  strata  adjacent  to 
the  Red  Sea  and  western  Indian  Ocean,  is  one  of 
ten  extant  genera  that  comprise  a  yiew  tribe. 
Decatopectinini. 

The  system  of  measurement  used  here  follows 
that  in  Waller  (1969,  1972),  in  which  shell  height 
is  measured  as  the  perpendicular  distance  be- 
tween the  active  outer  ligament  or  hingeline  and 
a  parallel  line  drawn  through  the  most  ventral 
point  on  the  ventral  margin.  References  to 
numbers  of  teeth  in  the  ctenolium  refer  only  to 
the  active  ctenolium  (Waller,  1984),  not  the  in- 
active ctenolium  partially  obscured  by  the 
growth  of  the  anterior  auricle.  The  umbonal 
angle  is  the  angle  between  lines  diverging  from 
the  origin  of  growth  and  passing  through  the 
estimated  points  of  union  between  the  free 
margins  of  the  auricles  and  the  disk  flanks. 
Observation  of  microsculpture  was  by  means  of 


a  Wild  M-.5  binocular  microscope  at  magnifica- 
tions not  exceeding  x  50.  Measurement 
employed  an  ocular  micrometer  as  well  as  a 
goniometer  eyepiece. 

The  term  antimarginal  is  introduced  with 
reference  to  microsculpture  that  maintains  a 
nearly  perpendicular  relationship  to  the  shell 
margin,  exemplified  by  the  Camptonectes  micro- 
sculptural  pattern  (illustrated  in  Waller,  1972). 
In  contrast,  sculptural  features  such  as  plicae, 
which  are  described  as  radial,  are  perpendicular 
to  the  shell  margins  only  in  the  midventral 
region  and  may  be  nearly  parallel  to  margins  in 
the  region  of  the  disk  flanks.  These  terms  com- 
plement the  term  commarginal,  which  has  come 
into  broad  usage  for  features  that  are  parallel  to 
shell  margins.  I  also  propose  new  terms  found  to 
be  useful  for  the  description  of  hinge  teeth  in  the 
Pectinacea.  In  the  right  valve,  resilial  teeth 
originate  near  the  origin  of  growth  of  the  shell 
and  approximately  parallel  the  anterior  and 
posterior  sides  of  the  resilium,  at  least  in  early 
ontogeny.  One  such  tooth  borders  each  side  of 
the  resilium,  as  in  Chlamys  islandica  (Miiller, 
1776).  Dorsal  teeth  lie  immediately  ventral  to 
the  outer  ligament,  one  such  tooth  bordering  the 
outer  ligament  on  each  side  of  the  hinge,  and 
may  originate  at  some  distance  from  the  origin 
of  growth  (arrows  in  Fig.  13).  Intermediate  teeth 
lie  between  resilial  and  dorsal  teeth,  are  com- 
monly differentially  developed  on  the  anterior 


40      THE  NAUTILUS 


April  :^0,  1986 


Vol.  100(2) 


and  posterior  sides  of  the  hinge,  and  may  be 
multiple,  as  in  Decatopecten  and  Pecten. 

The  anatomy  of  Snmaiipecfev  cranrnerorum- 
cannot  Ix'  described  at  {)resent,  because  no  soft 
parts  were  available  to  me. 

Systematics 

C/a.s.s  Bivalvia  Linnaeus,  1758 

Subclass  Pteriomorphia  Beurlen,  1944 

[e)n<')id..  Boss,  1982] 

Superorder  Eupteriomorphia  Boss,  1982 

Order  Ostreoida  Waller,  1978 

Suborder  Pectinina  Waller,  1978 

Superfamily  Pectinacea  Rafinesque,  1815 

[emend. .  Waller,  1978] 

Family  Pectinidae  Rafinesque,  1815 

[emend.,  Waller,  1978] 

Subfamily  Pectininae  Rafinesque,  1815 

Tribe  Decatopectinini,  new  tribe 

Diagnosis:  Pectinidae  having  very  closely 
spaced  commarginal  lamellae  at  least  in  early 
ontogeny  and  commonly  throughout  life,  the 
spacing  commonly  ranging  from  about  30  to  70 
lamellae  per  two-millimeter  distance  along  a 
radius  in  center  of  disk  at  height  of  10  mm;  infla- 
tion of  left  beak  very  low,  only  very  slightly  ex- 
ceeding that  of  right  beak,  the  left  beak  extend- 
ing only  very  slightly  dorsal  to  hingeline;  anti- 
marginal  microsculpture  very  fine  and 
restricted  to  early  ontogeny  of  disk  to  a  distal 
limit  slightly  beyond  origins  of  radial  plicae, 
absent  from  disk  flanks  and  auricles;  dentition 
dominated  by  dorsal  and/or  intermediate  teeth, 
the  latter  sometimes  multiple  or  sometimes 
absent;  resilial  teeth  low  or  absent. 

Type  Germs:  Decatopecten  Ruppel  in  Sowerl)v, 
1839. 

Taxonomic  Composition-The  new  tribe  con- 
tains the  following  extant  genera,  listed  with 
their  type  species,  geographic  region,  and  some 
common  synonyms: 

Anguipecten  Lall,  Bartsch,  and  Rehder,  193S, 
type  spac'w^  Anguipecten g7-egoryi  Dall,  Bartsch, 
and  Rehder,  1938  [junior  synonym  of  Perl  en 
Uintherti  Souverbie  //(  Sourverbie  and  Mon- 
trouzier,  1874],  tropical  Indo-Pacific. 

Annachlanii/s  Iredale.  1939,  type  species 
Pecten  Icopardus  Reeve,  1853  [junior  synonym 
o\  Pecten  flabellat us  Lamarck,   1819|.  tropical 


western  Pacific  and  eastern  Indian  Oceans. 

Bractechlamys  Iredale,  1939,  type  species 
Bractechkimys  evecta  Iredale,  1939  [junior 
synonym  of  Pecten  vexillum  Reeve,  1853], 
tropical  Indo-Pacific  and  Western  Atlantic. 

Decatopecten  Ruppel  in  Sowerby,  1839 
[senior  sync^nym  of  Comptopallium  Iredale, 
1939],  type  species  Ostrea  plica  Linnaeus,  1758, 
tropical  Indo-Pacific. 

Excellichlamys  Iredale,  1939,  type  species 
Pecten  spectabilis  Reeve,  1853,  tropical  Indo- 
Pacific. 

Flexopecten  Sacco,  1897  [senior  synonym  of 
Glabropecten  Sacco,  1897,  Lissopecten  Verrill, 
1897,  and  Proteopecten  Monterosato,  1899], 
type  species  Ostrea  jlexuosa  Poll,  1795,  Mediter- 
ranean and  adjacent  eastern  Atlantic. 

Gloripallium  Iredale,  1939,  type  species 
Ostrea  jialUitin  Linnaeus,  1758,  tropical  Indo- 
Pacific. 

Juxtamusiiim  Iredale,  1939,  type  species 
Juxtatnusiutn.  ublectatum  Iredale,  1939  [junior 
synonym  of  Pecten  (Chlamys)  coudeini  Bavay, 
1902],  tropical  western  Pacific  and  Indian 
Oceans. 

Mirapecten  Dall,  Bartsch,  and  Rehder,  1938, 
type  species  Mirapecten  thaanumi  Dall, 
Bartsch,  and  Rehder,  1938  [junior  synonym  of 
Pecten  mirificus  Reeve,  1853],  tropical  Indo- 
Pacific. 

Somalipecten  Waller,  new  geyius.  type  species 
Sonialipeeten  erannieroriun  Waller,  new  species 
described  herein,  tropical  western  Indian 
Ocean. 

St  rat  Igraphic  Range:  Paleocene  to  present. 

Discussion:  In  view  of  the  common  assump- 
tion that  commarginal  increments  in  many 
bivalves  are  periodic,  indicative  of  growth  rate, 
and  hence  subject  to  environmental  modifica- 
tion, it  would  seem  inapjiropriate  to  use  com- 
marginal spacing  as  one  of  the  prime  morpho- 
logical differentia  for  a  tribe.  However, 
evidence  is  accumulating  showing  that  growth 
increments  are  not  always  periodic  and  may 
occur  in  ways  that  are  taxonomically  specific 
and  hence  genetically  determined  (Jones,  1981; 
Ohno,  1985).  Recently,  Helm  and  Malouf  (1983) 
suggested  that  there  may  be  a  minimal  required 
distance  between  successive  commarginal 
ridges  in  the  Atlantic  Bay  Scallop.  .Argopecten 
irradians  (Lamarck,    1819),   and   that   this  re- 


Vol.  100(2) 


Ai)ril  30,  1986 


THE  NAUTILUS      41 


(luirenieiit  may  oN'erride  any  peiioiJicity  when 
the  rate  of  shell  growth  is  low.  (iruffydd  (1981) 
showed  that  in  Peden  nidxhitiis  (Linnaeus, 
17.'i8)  faster  growth  is  achieved  by  increasing 
both  the  width  of  growth  increments  and  the 
number  of  ridges  formed  per  unit  of  time. 
Whatever  the  cause  of  j)rojecting  commarginal 
lamellae  in  the  Pectinidae,  it  is  clear  that  these 
features  are  more  closely  spaced  in  members  of 
the  trilie  Decatopectinini  than  in  other  scallops 
at  a  comparable  shell  size  at  least  during  early 
ontogeny.  With  few  exceptions,  genera  outside 
the  Decatopectinini  have  fewer  than  25  com- 
marginal lamellae  per  two-millimeter  space  at  a 
height  of  10  mm,  and  many  of  these  have  as  few 
as  5  to  1.5. 

As  can  be  seen  from  the  above  list  of  geo- 
graphic ranges  of  genera,  the  Decato{:)ectinini 
are  largely  Indo-Pacific  at  present,  and  my  own 
unpublished  data  on  fossil  distriliutions  in- 
dicates that  the  trilie  has  been  primarily  Indo- 
Pacific  throughout  its  geologic  history.  The  two 
exceptional  genera  which  also  occur  in  the 
Atlantic,  Bracfixhiamifs  and  Flexopecten.  both 
appear  to  have  dispersed  into  the  Atlantic  from 
the  Indo-Pacific,  but  from  opposite  directions. 

The  relationship  of  Nodiperten  Dall,  1898,  and 
Lyropertev  Conrad,  1862,  to  the  Decatopectinini 
is  close,  but  both  of  these  genera  have  more- 
widely-spaced  commarginal  lamellae,  stronger 
intermediate  teeth,  and  coarser  umbonal  micro- 
sculpture.  A  full  account  of  the  morphological 
differences  that  distinguish  genera  in  the 
Decatopectinini,  as  well  as  a  discussion  of  fossil 
history,  zoogeography,  and  relationships  to 
other  suprageneric  units  within  the  Pectinidae, 
is  in  preparation. 

Somalipecten  Waller,  new  genus 
Type    Species:    Somalipecten    cranynerormn 
Waller,  new  species,  from  off  Somalia,  depth 
150  to  300  m. 

Diagnosis:  Plicate  Decatopectinini  having 
both  valves  convex,  byssal  notch  only  moderate- 
ly deep,  left  umbo  only  slightly  convex  or  flat- 
tened, and  disk  with  uneven  curvature,  incipient 
ledging,  or  widely  spaced  nodes  at  least  in  early 
ontogeny;  secondary  radial  costae  present  dis- 
tally;  auricular  costae  on  right  posterior  auricle 
weakly  developed  and  few  in  number  or  absent; 
enlarged  scales,  if  present,  limited  to  left  valve; 


dentition  dominated  l.)y  dorsal  teeth,  intermedi- 
ate teeth  weak,  resilial  teeth  commonly  absent. 
TaxoYiowic  Content:  The  new  genus  includes  a 
number  of  fossil  species  from  the  western 
Indian  Ocean  region  described  in  publications  by 
Cox  (1929)  and  Eames  and  Cox  (1956):  Chlamys 
(Acquipecten)  /(trsanensis  Cox,  1929,  C.  (A.) 
isthmica  (Fuchs,  1878),  C.  (A.)  leesi  Cox,  1929, 
C.  (A.)  lessepsi  (Fuchs,  1878),  C.  (A.)  pseudola 
Eames  and  Cox,  1956,  C.  (A.)  iverthi  (Philippi, 
1901),  and  C.  (A.)  wyllei  Cox,  1929. 

Sfrdtigrajihic  Range:  Upper  Miocene  to  pre- 
sent. All  of  the  fossil  species  listed  above,  with 
the  exception  of  S.  pseudola,  are  from  deposits 
adjacent  to  the  Red  Sea  (Egypt,  Sudan,  Saudi 
Arabia),  the  western  Indian  Ocean  (Somalia, 
Kenya,  Tanzania,  Zanzibar),  and  the  Arabian 
Sea  (southeastern  Saudi  Arabia,  Iran).  Their 
age  was  called  "post-Miocene"  by  Cox  (1929), 
and  some  are  probably  as  young  as  late  Pleisto- 
cene or  Holocene.  S.  pseudola  from  Iran  was 
said  by  Eames  and  Cox  (1956)  to  range  from 
Upper  Miocene  to  Pliocene.  Somalipecten  cran- 
merorum.  new  species,  is  the  only  known  living 
species. 

Comparison:  A  somewhat  flattened  left  umbo 
is  also  present  in  Annachlamys,  which  differs 
from  Somalipecten  in  having  a  wider  umbonal 
angle,  in  having  commarginal  lamellae  which 
revert  to  a  far-set  condition  late  in  ontogeny, 
and  in  lacking  nodes,  enlarged  scales,  or  exten- 
sive secondary  radial  costae.  Some  members  of 
Bractechlamys.  specifically  B.  langfordi  (Dall, 
Bartsch,  and  Rehder,  1938)  and  B.  noduliferum 
(Sowerby,  1842),  have  nodes  and  ledges  but  dif- 
fer from  Somalipecten  in  having  persistent  deep 
byssal  notches,  strong  intermediate  teeth,  and 
strong  costae  on  all  auricles.  Y^xi'dnt  N odipecten 
Dall,  1898,  and  fossil  Lyropecten  Conrad,  1862, 
and  Macrochlamis  Sacco,  1897,  differ  in  having 
far-set  commarginal  lamellae  and  much 
stronger  intermediate  hinge  teeth.  Flexopecten 
of  the  Mediterranean  and  eastern  Atlantic  dif- 
fers in  hinge  details,  the  right  dorsal  teeth  being 
much  weaker  or  absent,  and  lacks  nodes  or 
enlarged  scales.  Mirapecten  maintains  a  deep 
byssal  notch  throughout  ontogeny  and  has 
enlarged  scales  on  at  least  the  posterior  plica  of 
both  valves.  Notochlamys  Cotton,  1930,  differs 
from  Somalipecten  in  having  persistent 
shagreen  microsculpture,  and  Mesopeplum  dif- 


42      THE  NAUTILUS 


A()ril  30,  1986 


Vol.  100(2) 


fees  in  having  far-set  coinmarginal  lamellae  and 
prominent  resilial  teeth. 

Somalipecten  cranmerorum 

Waller,  ncir  spceivs 
Figs.  M.3 
Diagnoda:  Somalipeden  having  four  major 
pHcae  on  right  valve  and  three  on  left,  the  left 
valve  also  having  a  single  plica  of  smaller  ampli- 
tude adjacent  to  each  disk  flank;  enlarged  distal- 
l.y  concave  or  enclosed  scales  few  in  number  and 
widely  spaced,  limited  to  tops  of  plicae  of  left 
valve. 

Description:  Disk  Outline -Moderately  large, 
with  height  commonly  40  to  70  mm  and  seldom 
exceeding  length,  the  ratio  of  height  to  length 
commonly  0.88  to  1.01;  outline  acHne  or  slightly 
prosocline,  rarely  slightly  opisthocline,  and 
equilateral;  anterior  and  posterior  extremities 
of  disk  narrowly  rounded,  ventral  margin  broad- 
ly rounded;  umbonal  angle  ranging  from  93  to 
104°;  both  valves  convex,  the  right  more  so  than 
the  left,  the  umbone  of  left  valve  flattened;  ratio 
of  convexity  of  closed  valves  to  height  0.22  to 
0.37,  averaging  0.31. 

Auricle  Outlines -Right  anterior  auricle  with 
byssal  notch  only  moderately  deep,  bordered  on 
its  ventral  side  by  an  active  ctenolium  of  from  3 
to  5  delicate,  closely  spaced  teeth,  which  may  be 
obsolescent  in  largest  specimens;  other  auricles 
pointed,  their  free  margins  forming  acute 
angles  with  hinge  line;  anterior  auricles  ex- 
ceeding posterior  in  length,  the  ratio  of  length 
of  anterior  outer  ligament  to  length  of  posterior 
outer  ligament  commonly  1.04  to  1.24;  ventral 
migration  of  ligament  system  absent  even  in 
largest  individuals. 

Exterior  Shell  Surface -Right  disk  with  4 
major  plicae,  at  least  the  central  ones  beginning 
at  a  shell  height  of  between  2  and  3  mm  as  pairs 
of  low  rounded  costae  with  narrow  raised  crests, 
each  pair  then  merging  into  a  single  broad 
rounded  plica  at  shell  height  of  10  to  15  mm,  the 
broad  plica  becoming  flattened  and  bifid  with 
the  introduction  of  a  median  groove  at  a  shell 
height  of  about  35  to  40  mm;  anteriormost 
major  plica  bordered  anteriorly  in  early  onto- 
geny by  a  single  costa  along  edge  of  disk  flank; 
posteriormost  major  plica  bordered  posteriorly 
by  a  pair  of  costae  in  early  ontogeny.  Left  disk 
with  three  major  plicae  bordered  on  anterior 


and  posterior  by  a  single  lower  plica,  the  broad 
interspaces  broken  by  introduction  of  smaller 
median  plicae  corresponding  to  the  median 
grooves  on  right  plicae.  Secondary  radial  costae 
beginning  on  tops  of  right  plicae  at  shell  heights 
between  4  and  12  mm  and  across  both  plicae  and 
interspaces  of  both  valves  by  a  height  of  40  to  55 
mm.  Disk  flanks  without  radial  or  antimarginal 
costae.  Right  anterior  auricle  commonly  with  5 
to  7  costae  at  margin;  other  auricles  with  highly 
variable  costation,  the  right  posterior  auricle 
commonly  lacking  or  having  only  a  few  costae. 
Enlarged,  distally  concave  scales,  sometimes 
closed  to  form  hollow  knobs,  limited  to  tops  of 
the  five  major  plicae  of  left  valve,  few  in  number 
and  widely  spaced,  commonly  fewer  than  5  per 
plica,  rarely  absent  or  as  many  as  8  per  plica. 
One  or  two  commarginal  ledges  commonly  pre- 
sent in  distal  fourth  of  disk.  Prismatic  stage  of 
right  valve  extending  to  shell  height  of  1.8  to  2.3 
mm.  Microsculpture  on  left  beak  and  early  post- 
prismatic  stage  of  right  valve  before  start  of 
commarginal  lamellae  consisting  of  exceedingly 
fine  antimarginal  striae.  Commarginal  lamellae 
closely  spaced  throughout  ontogeny,  first  ap- 
pearing in  interspaces  of  right  disk  at  shell 
heights  between  4  and  6  mm. 

Internal  Features- Dentition  with  dorsal 
teeth  dominant,  intermediate  teeth  very  weak 
or  absent,  resilial  teeth  absent.  Single  crus  pre- 
sent on  inner  surface  of  shell  beneath  each  disk 
tlank,  a  second  crus  sometimes  present  on  ven- 
tral fourth  of  posterior  auricles;  other  crura  or 
marginal  denticles  generally  absent  on  inner 
surfaces  of  auricles.  Inner  surfaces  of  plicae 
with  carinate  edges  and  commonly  with  numer- 
ous radially  elongate  marginal  denticles. 
Ostracum  inside  pallial  line  consisting  entirely 
of  lathic  calcite  with  coarse  irregular  patches  of 
folia;  crossed  lamellar  aragonite  absent  in 
mature  shells. 

Color -Basal  pigment  red,  orange-red,  or 
orange,  either  solid  or  broken  by  exceedingly 
fine  white  mottling  and/or  by  bold  oblique  bars 
or  chevrons  of  white;  dark  pigment  commonly 
present  on  ventral  sides  of  enlarged  scales.  Pig- 
ment more  subdued  on  right  valve  than  on  left, 
the  bold  patterns  generally  restricted  to  left 
valve.  Beaks  commonly  with  fine  intersecting 
diagonal  white  lines.  Interior  of  shell  commonly 
{)igmented  outside  pallial  line  and  more  rarely 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      43 


FIGS.  1-9.  Holotype  of  Somali  jjecten  cranmerorum  Waller,  new  genus  and  species,  USNM  8590.34.  length  .53.8  mm,  from  off 
Somalia,  150-300  m.  1-6,  Left  exterior,  dorsal,  right  exterior,  anterior,  left  interior,  and  right  interior  views,  ammonium 
chloride  coating.  7,  Left  exterior,  uncoated.  8,  Detail  of  commarginal  lamellae  in  central  interspace  of  right  valve  at  heights 
between  14.7  and  17.1  mm,  coated  with  ammonium  chloride.  9.  Right  exterior,  uncoated. 


44      THE  NAUTILUS 


April  30,  1986 


Vol.  100  (2) 


FI(;S.  10-13.  Views  of  two  paratypes  o{ Somalipectcn  cranmerorum  Waller,  new  genus  and  species,  coated  with  ammonium 
chloride.  10,  Right  exterior  of  the  largest  paratype,  USNM  859035,  length  80.5  mm,  from  off  Somalia,  150-300  m.  11,  Closed 
scale  with  artificial  perforation  and  an  open  scale  on  a  central  plica  of  left  valve  of  another  paratype  returned  to  Mr.  Dan,  width 
of  open  scale  4.8  mm,  shell  height  at  level  of  open  scale  57  mm.  12-13,  Left  exterior  and  right  hinge  area  of  the  largest 
paratype,  USNM  835095,  shell  length  80.5  mm,  hinge  length  37  mm.,  arrows  in  Fig.  13  point  to  dorsal  hinge  teeth. 


inside  jiallial  line  by  the  same  color  present  on 
exterior. 

Comparison:  Somalipecten  cranmerorum, 
new  species,  differs  from  all  of  the  fossil  species 
mentioned  in  the  above  discussion  of  Somali- 
pecten, new  genus,  in  having  fewer  major  plicae. 
The  smallest  number  of  plicae  among  the  fossil 
taxa  occurs  in  Sovialipecfen  wyllpi  (Cox,  1929), 
which  has  an  additional  pair  of  plicae  on  each 
valve,  stronger  intermediate  hinge  teeth,  and  no 
enlarged  scales  on  the  left  valve.  S.  cranmero- 
rum superficially  resembles  the  more  coarsely 
plicate  varieties  of  Flexopecten  ,(//(;/>cr  (Linnaeus, 
1758)  living  in  the  Mediterranean,  Init  that 
species  lacks  nodes  and  enlarged  scales  and  dif- 
fers in  hinge  details,  as  discussed  above  in  the 
comparison  of  genera.  None  of  the  extant  or 
extinct  species  of  Nodipectcn  have  as  closely 
spaced  commarginal  lamellae,  and  they  have 
much  more  massively  developed  dorsal  and 
intermediate  hinge  teeth. 

Ecdiogy:  The  habitat  of  the  new  species  is 
known  only  from  the  data  provided  by 
Taiwanese  fishermen,  who  said  that  the  shells 
were  trawled  off  Somalia  at  depths  of  150  to  .'500 
m.  Because  they  have  not  divulged  the  exact 


locality,  it  is  not  known  whether  the  specimens 
are  from  the  Gulf  of  Aden  or  the  Indian  Ocean 
side  of  Somalia.  Mr.  T.  C.  Lan  of  Taipei, 
Taiwan,  who  obtained  the  specimens  from  the 
Taiwanese  fishermen,  has  provided  a  list  of 
associated  species,  some  of  which  would  appear 
to  shed  light  on  the  locality  of  the  pectinids. 
Strombu>!  otdi  Emerson,  1965,  was  originally 
described  from  the  Indian  Ocean  side  of  Somalia 
in  the  vicinity  of  Obhia  and  Mogadiscio  (Emer- 
son, 1965),  and  recently  it  also  has  been  found 
off  Oman  in  the  Arabian  Sea  (David  Hargreave, 
/)cr.s-.  com..  1986).  It  is  not  known  to  occur  in  the 
Red  Sea  or  the  Gulf  of  Aden.  Cypraea  hroderipi 
Soweri)y,  1832,  Cypraea  marginalis  Dillwyn, 
1SI7,  and  Vanum  Iruncafum  (Sowerby,  1892) 
are  known  mainly  from  the  Indian  Ocean  and 
not  the  Red  Sea  (Abbott  and  Dance,  1982),  and 
Mi))><ichlamys  toicn.scndi  (G.  B.  Sowerby  III, 
1.S95)  is  known  mainly  from  the  Arabian  Sea  off 
Pakistan  and  from  the  Gulf  of  Oman.  Although 
these  data  suggest  that  S.  cranmerorum  is  from 
the  Indian  Ocean  side  of  Somalia,  the  fossil 
species  that  is  morphologically  the  closest,  S. 
iryllfi  (Cox,  1929),  is  from  a  raised  beach  on  the 
Red  Sea  coast  in  Sudan. 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      45 


The  epifauna  of  the  shells  ofS.  cranmerorum 
gives  some  clues  to  living  habits.  The  lower 
(right)  valves  of  many  of  the  uncleaned  speci- 
mens are  encrusted  by  cheilstome  bryozoans  at 
least  in  the  dorsal  region  and  in  some  cases  ex- 
tensively over  the  valve.  This  indicates  that  the 
lower  valve  was  not  buried  in  fine  sediment  and 
that  the  scallop  must  have  lived  attached  by  a 
byssus  above  the  bottom  or  on  a  hardground 
bottom.  The  presence  of  calcareous  algae  on  the 
upper  valve  suggests  that  the  specimens  lived  in 
the  photic  zone,  as  does  also  their  association 
with  algal  grazers  such  as  Strombus. 

Thirteen  of  the  52  specimens  of  S.  cranmero- 
rum examined  have  circular  corroded  patches 
within  which  a  borehole  occurs  over  the  antero- 
dorsal  disk  flank  over  the  approximate  position 
of  the  scallop's  mouth.  These  features  are  iden- 
tical to  the  scars  produced  by  the  calyptreacean 
gastropod  Capulus  danieli  (Crosse,  1858) 
described  by  Orr  (1962;  see  also  Matsukuma, 
1978)  from  another  Indo-Pacific  member  of  the 
Decatopectinini,  Bractechlamys  vexillum 
(Reeve,  1853).  On  the  basis  of  gut  contents, 
absence  of  damage  to  soft  parts  other  than  the 
mantle  through  which  the  boring  passes,  and 
evidence  of  repair  and  continued  survival,  Orr 
concluded  that  the  snail-scallop  relationship  is 
one  of  antagonistic  symbiosis,  not  true  para- 
sitism, the  snail  stealing  food,  probably  in  mucus 
strings,  from  the  mouth  region  of  the  scallop. 
On  all  but  one  of  the  scarred  and  bored  scallops, 
the  shell  damage  is  on  the  left  umbo.  The  fact 
that  gastropod  was  able  to  survive  on  the  lower 
valve  is  additional  evidence  that  the  scallop  lives 
attached  above  the  sediment  surface. 

Etymology:  This  species  is  named  in  honor  of 
Roberta  D.  Cranmer  and  her  late  husband, 
Charles  E.  Cranmer,  of  Louisville,  Kentucky, 
whose  personal  involvement  and  quiet  philan- 
thropy have  greatly  benefited  many  people  and 
organizations,  a  number  of  which  have  made 
significant  contributions  to  malacology. 

Holotype:  USNM  859034,  a  pair  of  matching 
valves,  height  50.0  mm,  length  53.8  mm,  con- 
vexity across  closed  valves,  15.5  mm,  collected 
by  Taiwanese  fishermen  off  Somalia  at  a  depth 
between  150  and  300  m. 

Material:  In  addition  to  the  holotype,  USNM 
859034,  51  paratypes  were  studied,  all  paired 
valves  from  the  same  locality  at  150-300  meters 


off  Somalia.  Twelve  of  these  paratypes  are 
deposited  in  the  U.S.  National  Museum  of 
Natural  History  under  the  catalogue  numbers 

859035  (the  paratype  illustrated  herein)  and 

859036  (eleven  unillustrated  paratypes).  Thirty- 
one  of  the  remaining  paratypes  were  returned 
to  Mr.  Donald  Dan,  and  one  paratype  was  sent 
to  each  of  the  following  eight  museums: 
American  Museum  of  Natural  History,  New 
York;  Natural  History  Museum  of  Los  Angeles 
County,  Los  Angeles;  British  Museum  (Natural 
History),  London;  Museum  National  d'Histoire 
Naturelle,  Paris;  Rijks-museum  van  Naturlijke 
Historie,  Leiden;  Australian  Museum,  Sydney; 
Western  Australian  Museum,  Perth;  and  Na- 
tional Science  Museum.  Tokyo. 

Acknowledgments 

The  possibility  that  the  scallop  shells  from  off 
Somalia  were  of  a  new  species  was  first  con- 
sidered by  Mr.  Donald  Dan  of  Maryland  and  Mr. 
Al  Demartino  of  Naples,  Florida,  who  brought 
specimens  to  my  attention.  I  am  particularly 
grateful  to  Mr.  Dan  for  providing  a  large 
number  of  specimens  for  suiy,  for  under- 
writing the  cost  of  donating  sot-  of  these  to  the 
U.S.  National  Museum  and  othe  •  nuseums,  and 
for  providing  encouragement  id  numerous 
helpful  suggestions.  Mr.  Shin  Rong  Hwang  of 
Taipei  supplied  Mr.  Dan  with  the  specimens  ob- 
tained from  the  fishing-boat  captain,  and  Mr.  T. 
C.  Lan  of  Taipei  provided  information  on 
associated  species.  Mr.  C.  Patrick  Nuttall  and 
Dr.  Noel  Morris  kindly  provided  access  to  and 
assistance  with  the  fossil  collections  in  the 
British  Museum  (Natural  History).  Drs.  Harald 
Rehder  and  Richard  S.  Houbrick,  Department 
of  Invertebrate  Zoology,  Smithsonian  Institu- 
tion, and  Dr.  John  Pojeta,  Jr.,  U.S.  Geological 
Survey,  Washington,  D.('.,  kindly  reviewed  the 
manuscript  and  offered  suggestions  for  its  im- 
provement. My  assistant,  Mr.  Warren  C.  Blow, 
prepared  the  prints  and  plates. 

LITERATURE  CITED 

Abbott,  R.  T.  and  S.  P.  Dance.  1982.  Cvmperidnim  of  Sea- 
shells.  E.  P.  Button,  Inc.,  New  York,  ix  -i-  411  p. 

Boss,  K.  J.  1982.  Mojlusca,  p.  945-1166.  In  S.  P.  Parker, 
Synopsis  and  Classification  of  Living  Organisms,  Vol.  1. 
McGraw-Hill  Book  Co..  New  York. 

Cox.  L.  R.  1929.  Notes  on  the  Post-Miocene  Ostreidae  and 
Pectinidae  of  the  Red  Sea  Region,  with  Remarks  on  the 


46      THE  NAUTILUS 


Ai»ril  30,  198G 


100(2) 


Geological  Significance  of  their  l)istril>ulio[i.  I'mr. 
Malacol.  Sor.  London  18:lfir,-2()9.  pis.  11-13. 

Eames,  F.  E.,  and  L.  R.  Cox.  1950.  Some  Tertiary  Peclina- 
cea  from  East  Africa,  Persia,  and  the  Mediterranean 
Region.  Pror.  Malacol.  Sor.  London  32:1-68,  20  pis. 

Emerson,  W.  K.  1965.  Strombun  (Tricornis)  oldi  New 
Species.  Indo-Pncifir  MoUuxm  l(6):.397-398,  pi.  294. 

Criiffydd,  LI.  D.  1981.  Observations  on  the  Rate  of  Produc- 
tion of  External  Ridges  on  the  Shell  o{ Pecten  marimi/a  in 
the  Laboratory.  Jour.  Mnrinf  Biol.  Aax.  U.K.  61:401-4 1 1 . 

Helm.  N.  E..  and  R.  E.  Malouf.  1983.  Rate  of  Production  of 
External  Ridges  in  the  Bay  Scallop,  .4  n/opcc^'H  irradtmix. 
{.■\bstract)  Amer.  Zoologist  23(4):1024. 

Jones,  D.  S.  1981.  Repeating  Layers  in  the  Molluscan  Shell 
Are  Not  Always  Periodic.  Jour.  Pateo.  55(5):1076-1082. 

Matsukuma,  Akihiko.  1978.  Fossil  Boreholes  Made  by  Shell- 
Boring  Predators  or  Commensals.  I.  Boreholes  of  Capulid 
Gastropods.  Jupnnexe  Jour.  Molncol.  lVpnu.-<)  37(l):29-45. 

Ohno,  Terufumi,  1985.  Experimentelle  Analysen  zur 
Rhythmik  des  Schalenwachstums  einiger  Bivalven  und 
ihre    Palaobiologische     Bedeutung.     (Experimental 


Analyses  Concerning  the  Rhythm  of  Shell  Growth  in  some 

r^ivalves  and  its  Paleobiological  Implications).  Pnldconlo- 

unipliira.  Abt.  A,  289:63-123,  7  pis. 
Orr,  Virginia.  1962.  The  Drilling  Habit  or  ('apulux  daniM 

(Crosse)  (Mollusea:  Gastropoda).  The  Vrliger  5(2):63-67, 

pi.  7. 
Waller,  T.  R.  1969.  The  Evolution  of  the  Argopi'rtcn  gibbux 

Stock  (Mollusea:  Bivalvia).  with  Emphasis  on  the  Tertiary 

and    Quaternary    Species    of   Eastern    North    America. 

Polf(„il.  Sor.  Mfw.  3  (Jour.  Palco..  43(5),  suppl.),  125  p., 

7  pis. 
1972.  The  Pectinidae  (Mollusea:   Bivalvia)  of 

Eniwetok   Atoll,    Marshall    Islands.    The   VcIkjct   14(3): 

221-264,  8  pis. 
1978.  Morphology,  Morphoclines  and  a  New 

Classification  of  the  Pteriomorphia  (Mollusea:  Bivalvia). 

Phil.  Trans.  Royal  Soc.  London.  B,  284:345-365. 

_.  1984.  The  Ctenolium  of  Scallop  Shells:  Func- 


tional Morphology  and  Evolution  of  a  Key  Family-level 
Character  in  the  Pectinacea  (Mollusea:  Bivalvia).  Malarolo 
ijia  25(1):203-219. 


A  NEW  SPECIES  OF  TRITONIA  (NUDIBRANCHIA) 
FROM  SOUTHERN  CALIFORNIA  AND  BAJA  CALIFORNIA 

Hans  Bertsch' 

Biological  Sciences,  National  University,  Los  Angeles,  CA 

and 

Antonio  Mozqueira  Osuna 

Escuela  Superior  de  Acuacultura,  Centro  de  Estudios  Superiores 
del  Estado  de  Sonora  (CESUES),  Navojoa,  Sonora,  Mexico 

ABSTRACT 
Tritonia  myrakeenae  is  descrihed  as  a  new  species  from  the  Califomian  marine 
famial  province  and  is  ilisfinguished  anatomically  froni  the  north  Pacific  and 
north  Atlantic  species  q/" Tritonia. 


Although  the  opisthohranch  gastropod  fauna 
of  the  northeastern  Pacific  has  been  mono- 
graphed extensively  by  numerous  authors  (e.g., 
recently  by  Marcus,  19(il  a;  MacFarland,  19(i(v, 
Keen,  1971;  McDonald,  1983;  et  at.),  there  are 
still  numerous  new  distributional  records  (e.g., 
Bertsch,  1981;  Behrens,  1982;  Gosliner  & 
Millen,  1984)  and  new  species  descriptions  (e.g.. 


'Mailing  address:  6056  Beeman  Ave..  No.  Hollywood,  CA 
91606 


Gosliner,  1981;  Behrens,  1984;  and  Millen,  1985) 
being  published.  These  add  significantly  to  the 
known  species  of  opisthobranchs  that  are 
endemic  to  this  coastline  or  its  various  marine 
faunal  provinces,  or  that  are  shared  with  other 
marine  zoogeographic  regions.  In  this  paper  we 
describe  a  new  species  of  Tritonia  that  occurs  in 
the  Californian  marine  faunal  province;  the  ex- 
istence of  this  species  was  first  reported  over  5 
years  ago  (Behrens,  1980). 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      47 


Order  Nudibranchia 
Suborder  Dendronotacea 

Family  Tritoniidae  Lamarck,  1809 

Tritonia  myrakeenae  Bertsch  &  Mozqueira, 
n.  up. 

Syno)ii/>ny:  Tritonia  sp.  Behrens,  1980: 
102-103  (includes  a  color  photograph  of  the  liv- 
ing animal). 

Type  Locality:  Southeast  end  of  Isla  Cedros, 
Baja  California,  Mexico,  rocky  reef  west  of 
lighthouse,  in  Bahfa  Sudeste:  28°2'30"N; 
115°12'W. 

Type  Material:  Holotype:  Approximately  7 
mm  long  while  alive,  4.5  mm  preserved;  inter- 
tidal  zone,  rocky  reef  SE  end  of  Isla  Cedros;  leg. 
H.  Bertsch  and  Soa  Tsung,  26  December  1985. 
Deposited  in  the  collection  of  the  Los  Angeles 
County  Museum  of  Natural  History,  Dept.  of 
Malacology,  No.  LACM  2135. 

Paratype:  Approximately  6  mm  long  while 
alive,  4  mm  preserved;  intertidal  zone,  rocky 
reef  SE  end  of  Isla  Cedros;  leg.  H.  Bertsch  and 
Soa  Tsung,  26  December  1985.  Deposited  in  the 
collection  of  Los  Angeles  County  Museum  of 
Natural  History.  Dept.  of  Malacology,  No. 
LACM  2136. 

Para  types:  Three  specimens,  3.5,  4,  and  4.5 
mm  preserved  lengths;  intertidal  zone  in  front 
of  Hotel  Puesta  del  Sol,  El  Sauzal,  approximate- 
ly 8  km  north  of  Ensenada,  Baja  California, 
Mexico  (3r52'N;  116°41'W);  leg.  A.  Mozqueira, 
September  1984.  Deposited  in  the  collections  of 
California  Academy  of  Sciences,  Dept.  of  Inver- 
tebrate Zoology,  No.  CASIZ  061379. 

Additional  Records  and  Range:  The  authors 
and  Jesus  Pineda  have  found  specimens  infre- 
quently in  the  intertidal  region  at  El  Sauzal  (in 
front  of  Hotel  Puesta  del  Sol),  Baja  California, 
during  June,  September,  October  and  Novem- 
ber 1983,  and  February  and  March  1984. 
Behrens  (1980:  102)  reported  the  occurrence  of 
Tritonia  myrakeenae  (as  Tritonia  sp.)  from 
"Santa  Barbara  to  San  Diego,  California."  All 
known  records  of  this  species  are  only  from  the 
intertidal  region.  At  this  time,  the  known  r'ange 
of  Tritonia  myrakeenae  is  from  Santa  Barbara, 
southern  California.  U.S.A.,  to  the  SE  tip  of  Isla 
Cedros,  near  the  southern  limit  of  the  state  of 
Baja  California,  Mexico. 

Description:     A     thin,     elongate,     delicate- 


appearing  tritoniid,  6-18  mm  in  total  length 
(alive).  Oral  veil  with  4-7  (rarely  branching)  ten- 
tacular processes.  Foot  broadly  expanded 
laterally  beyond  sides  of  body;  anterior  margin 
gently  rounded,  bilabiate;  posteriorly  extending 
out  flat  past  the  body,  terminating  in  a  pointed 
tip.  Retractile  rhinophores  surrounded  by  basal 
sheath;  the  rhinophore  stalk  bears  10-22  vertical 
processes  which  encircle  the  shaft  below  the  tip. 
Dorso-lateral  edge  with  a  delicate,  undulating 
free  margin  from  which  grow  the  branchial  pro- 
cesses (often  9  on  each  side).  Each  branchial  ap- 
pendage branches  distally  into  3-5  filaments. 

Color  a  very  pale  dirty  orange-brown  (see 
color  photo  in  Behrens,  1980:  103);  oral  veil  and 
sides  of  foot  clear  of  color,  partly  opaque.  Dis- 
tinctive white  patches  occur  on  the  dorsum 
(often  as  a  matched  pair  on  each  side  of  the 
midline);  Behrens  (1980:  103)  illustrates  an 
animal  with  4  transverse  groups  of  opaque  white 
patches  (or  streaks)  arranged  down  the  length 
of  the  back.  Specimens  from  Ensenada  and  Isla 
Cedros  (Figiu'e  1)  had  fewer  white  splotches 
(usually  just  a  pair  located  about  Vs  of  the  body 
length  distant  from  the  anterior  end). 

Radula  (Figure  2 A)  about  1.1  mm  long,  0.65 
mm  across  at  widest  portion.  Radular  formulae 
of  2  specimens  collected  26  September  1983  at 
El  Sauzal  were:  22  (10.1.1.1.10)  and  25 
(8-10.1.1.1.8-10)  (widest  row  damaged  but  ap- 
peared to  have  12  outer  lateral  teeth).  Central 
rachidian  tooth  prc.iminent  (a|)proximately  63 
microns  wide),  with  3  posteriorly-directed  cusps 
(Figures  2B  and  3B).  First  lateral  tooth  well  dif- 
ferentiated from  the  others,  its  structure  a 
hooklike  cusp  curling  over  the  side  of  the  rachi- 
dian (Figiu'es  2B  and  3B).  Succeeding  laterals 
all  fairly  straight,  lilade-like  cusps. 

Jaws  elongate  (P^igin-e  3A),  each  about  1.3  mm 
long  and  0.37  mm  wide;  a  row  of  pointed  mam- 


5  mm 


FlCi.    1.   Tntiiiiin  iiiijrakfi'xiif.  xpt'c.  tii)i\,  dorsal  view  of  liv- 
ing animal.  Drawing  by  C.  Yanes. 


48      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


2  A 


2  B 


FIG.  2.  Camera  lucida  drawings  (scale 
represents  50(j)  of:  2A.  Major  shape  and 
features  of  entire  radula,  and  2B.  Raehi- 
dian  tooth,  innermost  lateral,  and  four 
outer  lateral  teeth  of  Tri.toiiia  niyra- 
keenae. 


FIG.  3.  Tritonia  myrakeenae.  sketches 
of  jaws  (3A)  and  selected  radular  teeth 
(3R,  rachidian,  innermost  lateral  tooth, 
and  two  outer  lateral  teeth). 


millate  denticles  begins  on  the  inner  margin  at 
the  joined  region  of  the  jaws,  continuing  as  a 
row  along  the  entire  inner  edge,  increasing 
about  Va  of  the  length  to  3-5  rows;  the  inner  den- 
ticled  margin  terminates  distally  in  a  small  blunt 
process  or  free  flange. 

Discusfiion:  Tritonia  myrakeenae  is  readily 
distinguished  from  other  north  Pacific  species  of 
Tritonia.  Tritonia  hawaiiensis  Pease,  1860, 
has  much  more  prominent  and  more  highly 
branched  branchial  processes  and  tints  of 
orange-pink  and  dirty  light-purple  (Bertsch  & 
Johnson,  1981:  84-85)  which  are  absent  in  our 
new  species.  Tritonia  insulae  (Baba,  1955)  has 


vermilion-tinted  gills  and  a  different  radular  for- 
mula: 30  (40-46.1.1.1.40-46).  Tritonia  diomedea 
Bergh,  1894,  has  a  distinct  white  line  bordering 
the  edge  of  the  foot,  and  a  larger  radular  for- 
mula. Tritonia  festiva  (Stearns,  1873)  has  a 
series  of  white  lines  and  loops  reticulating  on 
the  dorsum  and  a  radula  with  more  teeth  in  each 
row.  Tritonia  pickensi  Marcus  &  Marcus,  1967, 
has  a  distinct  white  swath  running  down  the 
center  of  the  dorsum  with  lateral  extensions  to 
the  branchial  processes;  although  it  has  a  similar 
radular  formula,  the  shape  of  the  rachidian 
tooth  (narrower  and  deeper  indentation  be- 
tween the  cusps)  and  the  presence  of  small  den- 
ticles on  the  cusp  of  the  first  lateral  tooth 
(Bertsch  &  Gosliner,  1984)  are  differentiating 
features. 

Species  of  Tritonia  in  the  North  Atlantic  are 
also  easily  distinguished  from  T.  myrakeenae. 
The  tropical  T.  welhi  Marcus,  1961  (b),  is  pink 
and  white,  and  the  shapes  of  the  teeth  are  differ- 
ent (cf.  Marcus  &  Marcus,  1967:  100,  fig.  130), 
and  T.  bayeri  Marcus  &  Marcus,  1967,  has  an 
opaque  white  diffuse  network  over  its  dorsum. 
Tritonia  nilsodhneri  Marcus,  1983,  is  rose-pink; 
T.  m.anicata  Deshayes,  1853,  has  red,  black  or 
olive-green  spots  on  the  dorsum;  T.  plebeia 
Johnston,  1828,  has  white  stippling  but  is  darker 
colored  with  brown  mottling  and  has  more  teeth 
per  half  row  (21-33);  T.  lineata  Alder  & 
Hancock,  1848,  has  2  conspicuous  longitudinal 
white  lines  running  lengthwise  down  the  dor- 
sum; although  T.  homhei-gi  Cuvier,  1803,  has 
white  splotches,  its  body  shape  is  different 
(tubercled  dorsum,  more  prominent  bilobed  oral 
veil)  and  the  radula  is  completely  different  (den- 
ticles on  teeth  of  young  specimens  and  27-158 
teeth  in  a  half  row).  Thompson  &  Brown  (1984) 
give  further  anatomical  characteristics  that 
distinguish  these  latter  5  species  from  T. 
myrakeenae.  Mrs.  Marcus  (1983)  details  the 
anatomy  of  other  Tritoniidae,  none  of  which  are 
comparable  with  Tritonia  myrakeenae. 

Etymology:  This  new  species  is  named  in 
honor  of  the  distinguished  malacologist,  Dr.  A. 
Myra  Keen  (1905-1986),  colleague  and  friend, 
who  included  nudibranchs  in  Sea  Shells  of 
Tropical  West  America,  second  edition  (this  was 
the  first  major  review  of  all  known  nudibranchs 
and  other  opisthobranchs  from  the  Panamic 
marine  faunal  province).  Her  persistent  scholar- 


Vol.  100(2 


Af)ril  30,  1986 


THE  NAUTILUS      49 


ship  and  encouragement  of  research  has  helped 
us  to  know  (and  hopefully  to  protect)  the 
mollusks  of  western  North  America. 

Acknowledgments 

Field  work  was  partially  supported  by  a  grant 
(to  the  senior  author)  from  Have  Mule  Will 
Travel  (Alpine,  California),  and  with  the  co- 
operation of  the  Escuela  Superior  de  Ciencias 
Marinas,  Universidad  Autonoma  de  Baja 
California,  Ensenada  (Guadalupe  Ballesteros, 
Director). 

We  especially  thank  Jesus  Pineda  for  collect- 
ing assistance  in  Ensenada,  and  the  Cedros 
Island  Have  Mule  Will  Travel  research  expedi- 
tion team  (including  Miss  Soa  Tsung)  of 
December  1985-January  1986. 

The  illustration  of  the  living  animal  of 
Tritonia  myrakeenae  (Figure  1)  was  drawn  by 
Senorita  Clara  Yanes. 


LITERATURE  CITED 

Behrens,  D.  W.  1980.  Pacific  coast  nudibranchs:  a  guide  to 
the  opisthobranchs  of  the  northeastern  Pacific.  Sea  Chal- 
lengers, Los  Osos.  112  pp. 

1982.   Sakuraeolis  en.osimensis  (Baba,    1930) 

(Nudibranchia:  Aeohdacea)  in  San  Francisco  Bay.  Veliger 
24(4):359-363. 

.  1984.  Notes  on  the  tergipedid  nudibranchs  of  the 


northeastern  Pacific  with  a  description  of  a  new  species. 
Veliger  27{l):6b-7l. 
Bertsch,  H.  1981.  Rectification  of  the  generic  placement  of 
Sclewdoris   tanyn   (Marcus,    1971).   comb.    tior..    a  nudi- 


branch  from  southern  California,  with  a  range  extension  to 
the  Gulf  of  California,  Mexico.  Veliger  23(3):217-220. 

Bertsch,  H.,  and  T.  Gosliner.  1984.  Tritoyiin  pickeiisi  (Nudi- 
branchia: Tritoniidae)  from  Baja  California,  Mexico.  Shells 
and  Sea  Life  16(9):138-139. 

Bertsch,  H.,  and  S.  Johnson.  1981.  Hawaiian  nudibranchs. 
Honolulu,  Oriental  Publ.  Co.,  112  pp. 

Gosliner,  T.  M.  1981 .  A  new  species  of  tergipedid  nudibranch 
from  the  coast  of  California.  Jour.  Moll.  Studies 
47:200-20,5. 

Gosliner,  T.  M.,  and  S.  V.  Millen.  1984.  Records  oiCuthoria 
pustulata  (Alder  &  Hancock,  1854)  from  the  Canadian 
Pacific.  Veliger  26(3):183-187. 

Keen,  A.  M.  1971.  Sea  shells  of  tropical  west  America,  sec- 
ond edition.  Stanford,  California.  Stanford  University 
Press,  1064  pp. 

MacFarland.  F.  M.  1966.  Studies  of  opisthobranchiate  mol- 
lusks of  the  Pacific  coast  of  North  .-Vmerica.  Mem.  Calif 
Acad.  Sci.  6:546  pp. 

Marcus.  Er.  1961  a.  Opisthobranch  mollusks  from  Califor- 
nia. Veliger  3  (Suppl.):l-85. 

1961  b.  Opisthobranchia  from  North  Carolina. 

Jour.  Elisha  Mitchell  Scient.  Soc.  77:141-151. 

Marcus,  Ev.  du  Bois  Reymond.  1983.  The  western  Atlantic 
Tritoniidae.  Bolm.  Zool.  Univ.  S.  Paulo  6:177-214. 

Marcus,  Er.  and  Marcus,  Ev.  1967.  American  Opisthobranch 
Mollusks.  Studies  Trop.  Oceanogr.  6:1-256. 

McDonald,  G.  1983.  A  review  of  the  nudibranchs  of  the  Cali- 
fornia coast.  Malacologia  24(1-21:114-276. 

Millen.  S.  V.  1985.  The  nudibranch  genera  Onchidons  and 
Diaphorodoris  (Mollusca,  Opisthobranchia)  in  the  north- 
eastern Pacific.  VV/((/fv28(l):80-93. 

Stearns,  R.  E.  C.  1873.  Descriptions  of  a  new  genus  and  two 
new  species  of  nudibranchiate  mollusks  from  the  coast  of 
California.  Proc.  Calif  Acad.  Sci.  5:77-78. 

Thompson,  T.  E.,  and  G.  H.  Brown.  1984.  Biology  of 
Opisthobranch  Molluacs.  Volume  II.  London,  The  Ray 
Society,  229  pp. 


UNIONIDAE  OF  THE  UPPER  CONNECTICUT  RIVER, 
A  VANISHING  RESOURCE' 

Arthur  H.  Clarke 

ECOSEARCH,  Inc.,  325  East  Bay  view 
Portland,  TX  78374 


Introduction 

It  was  my  good  fortune  to  meet  William  J. 
Clench  in  the  winter  of  1949-50,  shortly  after 
having  become  irrevocably  committed  to  shell 
collecting.  His  personality  was  so  enthusiastic 
and  friendly  and  the  Mollusk  Department  was  so 


'In  memory  of  my  good  friend  and  teacher,  William  J. 
Clench. 


impressive  that  I  soon  made  frequent  trips  to 
the  Museum  of  Comparative  Zoology  at  Har- 
vard, known  familiarly  as  the  "MCZ",  and  later 
became  a  Saturday  volunteer.  It  was  not  long 
after  that  I  resolved  to  seek  a  career  in 
malacology.  I  shall  always  be  grateful  to  Bill 
Clench  and  to  Ruth  Turner  for  the  inspiration  to 
make  that  decision  and  for  their  numerous  acts 
of  kindness  since  that  time. 


50      THE  NAUTILUS 


April  80,  1986 


Vol.  100(2) 


I  well  remember  our  first  collecting  trip  to  the 
Connecticut  River.  The  occasion  was  the  1952 
annual  field  trip  of  the  Boston  Malacological 
Club;  the  site  was  the  Connecticut  River  at 
South  Hadley,  Massachusetts;  the  leader  was 
Dr.  Clench;  the  weather  was  warm  and  sunny; 
and  the  water  was  low.  Bill,  as  his  friends  called 
him  with  affection  and  respect,  led  us  to  a  large 
sandbar  in  the  middle  of  the  river.  Beautiful 
mussels  were  everywhere  in  2  or  3  feet  of  water 
and  the  Lampsilis  cariosa  were  the  most 
massive  that  I  had  ever  seen. 

During"  the  following  years  Arthur  and  Louise 
Clarke  collected  widely  in  New  England  and  at 
several  localities  in  the  Connecticut  River 
System.  There  we  found  Alasmidontu  hetero- 
don.  atrangeAhismidonta  uruiulata.  and  several 
other  species  which,  to  my  neophyte  eyes,  were 
especially  marvelous.  Although  the  Connecticut 
River  fauna  had  recently  been  reported  upon 
(Clench  and  Russell,  1939,  1940),  it  still  merited 
additional  investigation.  It  was  not  until  nearly 
30  years  had  passed,  however,  that  I  was  able  to 
do  "it. 

It  is  appropriate  that  the  results  of  that 
survey,  although  principally  of  regional  interest, 
should  be  dedicated  to  Bill  Clench.  In  a  larger 
sense,  however,  all  of  my  research  has  been  in- 
fluenced by  his  teaching  and  it  will  always  con- 
tinue to  be  so.  I  would  also  like  to  thank  The 
Nature  Conservancy  for  funding  the  field  work 
in  1983  and  my  wife  Judith,  and  Drs.  L.  L. 
Master  and  T.  French,  for  valuable  field 
assistance.  Preparation  of  the  manuscript  was 
supported  by  ECOSEARCH,  Inc. 

Methods  and  Results 

The  primary  objective  of  the  1983  program 
was  to  determine  the  present  distribution  and 
abundance  of  Alasruidovfa  hetcrodon  (Lea)  in 
the  Connecticut  River  System  of  New  Hamp- 
shire and  Vermont.  During  the  course  of  that 
work  tabulation  of  habitat  attributes  and  of  the 
identity  and  abundance  of  all  other  mollusks  en- 
countered were  also  made.  The  detailed  results 
concerning/!,  keterodori  will  be  reported  as  part 
of  a  status  survey,  now  underway,  of  that 
species  throughout  its  entire  range,  but  the 
general  results  for  1983  are  reported  here. 

The  principle  field  activities  were  carried  out 
over   an    18-day   period    in    August  and   early 


September,  1983.  Collecting  equipment  in- 
cluded a  motor-assisted  canoe,  wet  suits,  view- 
ing boxes,  and  a  small  Kolkwitz  dredge.  A  total 
of  36  survey  areas  were  studied.  These  are 
shown  in  Map  1  and,  to  conserve  space,  are  ab- 
breviated and  listed  below  in  narrative  form. 

Station  List:  1,  Third  Conn.  L,  E  side,  N.H.;  2, 
Conn  R  between  Third  and  Second  Conn.  L;  3, 
Second  Conn.  L,  S  side;  4,  First  Conn.  L,  N  side, 
N.H.;  5,  Conn.  R  just  above  Francis  L,  N.H.;  6, 
Conn.  R  above  Canaan,  Vt.;  7,  Conn.  R  1.0  mi  N 
of  Tinkerville,  Columbia  Twp.,  N.H.;  8,  9,  Conn. 


Mass. 


FU;.   1.  The  upper  Connecticut  River  System  with  1983 
study  sites  indicated. 


Vol.  100  (:; 


April  30,  1986 


THE  NAUTILUS      51 


R,  by  canoe  from  N  Stratford  to  2.4  mi  above 
Stratford,  N.H.;  10,  Conn.  R,  2.4  mi  above 
Stratford,  N.H.;  11,  Conn.  R,  flooded  marginal 
pool,  above  bridge  at  Guildhall,  Vt.;  12-14, 
Conn.  R,  by  canoe  from  Comerford  Dam  to 
bridge  at  Barnet,  Vt.;  15,  Conn.  R,  S  Newbury, 
Vt.,  V4  mi  above  Oxbow  Bridge;  16,  Conn.  R,  1.0 
mi  NW  of  Piermont,  N.H.;  17,  Conn.  R,  Hart- 
land,  Vt.;  18,  Conn.  R,  Weathersfield  Bend,  4  mi 
S  of  Ascutney,  Vt.;  19,  Conn.  R,  below  bridge 
from  N  Walpole,  Vt.  to  Bellows  Falls,  N.H.;  20, 
Nulhegan  R,  0.5  mi  W  of  Bloomfield,  Vt.;  21, 
Paul  Stream,  1.1  mi  above  mouth,  Brunswick 
Twp.,  Vt.;  22,  Upper  Ammonoosuc  R,  Groves- 
ton,  N.H.;  23,  Forest  L,  Dalton  Twp.,  N.H.;  24, 
outlet  of  Hazen's  Pond,  2.5  mi  E  of  Whitefield, 
N.H.;  25,  lake  at  town  park,  Thetford  Twp.,  Vt; 
26  Mud  Pond,  Thetford  Center,  Vt.;  27,  Ompom- 
panoosuk  R,  0.5  mi  S  of  Union  Village,  Vt.;  28, 
Croydon  Branch,  1.0  mi  S  of  Croydon,  N.H.;  29, 
Sugar  R,  2  mi  SW  of  Croydon;  30,  Williams  R, 
0.5  mi  N  of  Rockingham,  Vt.;  31,  Ashuelot  Pond 
outlet,  3.0  mi  NE  of  Marlow,  N.H.;  32,  Ashuelot 
R,  0.4  mi  W  of  Ashuelot  Pond;  33,  Asheulot  R 
just  above  Marlow,  N.H.;  34,  Ashuelot  R,  be- 
tween Shaw's  Corner  and  Roundy's  Corner, 
N.H.;  35,  Ashuelot  R,  3.0  mi  S  of  Surry  Moun- 
tain Dam;  36,  Ashuelot  R,  2.4  mi  NW  of  center 
of  Keene,  N.H. 

In  most  cases  living  specimens  were  tabulated 
and  promptly  returned  to  their  habitats.  In  the 
tabulation  below,  each  station  number  is  fol- 
lowed by  a  syml)ol  which  reflects  the  number  or 
abundance  of  specimens  seen  there.  Numbers  of 
living  (no  parenthesis),  recently-living  (in  paren- 
thesis), and  subfossil  (in  parenthesis,  with  S) 
specimens  are  cited  separately.  Other  symbols 
are:  A  =  abundant  and  C  =  common.  No  mollusks 
were  found  at  stations  4,  5,  6,  12,  13,  14,  20,  22, 
24,  27-31,  33,  and  34. 

Species  List 
Unionidae.  M.  inargaritifera:  sta.  32,  (2S).  E. 
complnmita:  8,(1S);  15,3; 'l6,(25);  17,ca.500-i- 
(77);  18,2-H(1);  19.(5S);  23,12-h(l);  35,A;  36,A. 
An.  cataracta:  l,12-i-(6);  2,6-i-(2);  3,41 -i-(3); 
11,(1);  23,2-^(1).^/.  heterodon:  17,(5);  35,1-h(2); 
36,(1).  AL  undulata:  1,1 -(-(9);  3,2;  9,(1S);  17,(24); 
35,2-^(3);  36,4.  St.  undulata:  3,1;  17,(4);  35,1; 
36,3-f(l).  La7rip.  r.  radiata:  17,ca.300-i-(28); 
19,(4S). 


Sphaeriidae  Sphaerium  striafinuni:  17,11. 

Viviparidae.  Canipclonid  decisum:  10,3; 
16,27;  17,2;  36,18.  Cipanqopaludina  chinerhsis: 
25, A;  26,A. 

Lymnaeidae.  Juv.  lymnaeid,  sp.?:  7,  1. 

Physidae.  f^hi/.^n  heterostropha:  11, A;  16,1. 

Planorbidae.  Helisovia  anceps:  7,1;  8,1;  11,C; 
16,4;  17,4. 

Ancylidae.  Ferrisia  rivularis:  9, A;  18, A. 

Discussion  and  Conclusions 

According  to  Johnson  (1915),  Clench  and 
Russell  (1939,  1940),  and  Clarke  (1981),  and 
specimen-associated  data  in  the  MCZ,  USNM, 
and  ANSP,  10  species  of  freshwater  mussels  oc- 
curred in  the  Connecticut  River  prior  to  and 
during  part  of  the  first  few  decades  of  this  cen- 
tury. They  are  Elliptio  complanata,  Anodonta 
cataracta.  An.  implicatn.  Alasmidonta  hetero- 
don, Al.  undulata.  Al.  varicosa,  Strophitus 
undulatus,  Lampsilis  cariosa,  L.  ochracea  and 
L.  r.  radiata.  Three  of  these  (An.  implicata,  L. 
cariosa  and  L.  ochracea)  were  not  known  to 
ascend  the  river  above  Massachusetts.  Many 
widely-distributed  species  of  Sphaeriidae,  proso- 
branchs,  "and  pulmonates  were  also  known  to 
occur  there  (Johnson,  1915;  Clench  &  Russell, 
1939)  and  another  unionid  (Ligumia  nasuta)  is 
also  known  from  the  vicinity  of  Keene,  N.H., 
presumably  in  the  Ashuelot  River  drainage 
(Walker  and  Coolidge,  1908).  The  principle  ob- 
jective of  this  paper  is  to  discuss  the  unionids  of 
the  main  river,  but  the  records  listed  above  from 
some  tributaries  (e.g.  the  introduced  gastropod, 
Cipangopaludina  chinensis)  may  also  be  of 
interest. 

Early  in  the  course  of  our  work  it  became  ob- 
vious that  the  rnolluscan  fauna,  and  especially 
the  unionids,  of  the  Connecticut  River  has  been 
greatly  harmed  by  human  activities.  Dams,  the 
construction  of  which  began  in  the  1930's,  have 
altered  this  once  free-flowing  river  so  that  it  is 
now  chiefly  a  series  of  impoundments  within 
which  water  levels  are  drastically  manipulated 
by  computer  controlled  dams  for  the  generation 
of  electricity  and  for  flood  control.  Pollution 
from  the  pulp  and  paper  industry  has  further  ex- 
acerbated the  situation.  The  probable  distribu- 
tions of  the  unionids  in  the  upper  Connecticut 
River,  as  they  existed  prior  to  historical 
degredation  of  the  river,  are  shown  in  Figau-e  2. 


52      THE  NAUTILUS 


April  80,  1986 


Vol.  100(2) 


c 
< 


■D 

C 
3 


■D 

C 
3 


V 

(0 


4  5" 

P 

■ 

• 

44' 

• 

— 

4  3' 

• 

o 
•    o 

• 

• 

Vt/NH 

Mass 


FIG.  2.  Probable  distribution  of  unionids  in  upper  Connec- 
ticut River  prior  to  pollution  and  impoundment  (white  bars 
and  open  circles)  contrasted  with  their  1983  distributions 
(black  bars  and  black  spots).  Locations  are  expressed  in 
terms  of  north  latitude.  Pre-impoundment  distributions  are 
based  on  subfossil  specimens  (stations  8  and  19),  live 
specimens  collected  by  W.  .J.  Clench  from  Conn.  River  near 
M(mroe,  N.H.,  and  Wells  River,  Vt.,  in  1940,  other  records 
at  MCZ,  I'SNM,  and  ANSP,  and  Clench  and  Russell  (1939. 
1940). 


The  di.strihutioii  of  those  species  as  observed  in 
1983  are  also  shown  on  that  figure. 

Based  on  the  presence  or  absence  of  fresh- 
water mussels,  it  is  useful  to  consider  the  upper 
Connecticut  River  as  now  comprised  of  3  faunal 
regions.  Region  1  (stations  1-3)  includes  the 
uppermost  part  of  the  system,  i.e.,  the  Second 
and  Third  Connecticut  Lakes  and  the  interven- 
ing portion  of  the  Connecticut  River.  (The  u{)- 
permost  Fourth  Connecticut  Lake  is  small  and 
inaccessible  by  road;  it  was  not  sampled).  Signif- 
icantly, the  water  levels  in  this  part  of  the 
system  are  not  manipulated  but  are  left  to  fluc- 
tuate moderately  and  naturally.  This  region  con- 
tains large  poy)ulations  oi  Anodonta  rnturnrta. 
(specimens  of  which  have  a  i)eculiar  subarctic 
fades),  of  Alasmidontd  undulata  morph  sower- 
hyuna.  (an  ecophenotyj)e  which  has  been  shown 
(Clarke,  1981)  t<j  be  attributable  to  slow  growth 


in  cold,  hard-water  habitats)  and  a  small  popula- 
tion of  Strophitus  undulatus.  Elliptio  com- 
planata  does  not  occur  here. 

Region  2  (stations  4-14)  includes  the  area  from 
First  Connecticut  Lake  and  Lake  Francis  down- 
stream for  more  than  100  miles  to  near  the 
mouth  of  the  Ammonoosuc  River  in  Grafton  Co., 
N.H.  Although  one  apparently  fresh  pair  of 
valves  of  An.  cataracta  occurred  in  a  flooded 
pool  adjacent  to  the  Connecticut  River  (station 
11),  no  freshwater  mussels  occurred  alive  in  this 
whole  stretch  of  the  main  river.  Lakes  drained 
by  at  least  one  tributary  (Johns  River  in  Coos 
Co.,  N.H.)  do  contain  both  Elliptio  complnnata 
and  An.  cataracta,  however.  Water  levels  in 
First  Connecticut  Lake  and  in  Lake  Francis  are 
artificially  lowered  by  about  18  feet  during  the 
winter  to  provide  for  flood  control  and  water 
levels  in  the  Connecticut  River  throughout 
Region  2.  They  are  also  drastically  manipulated 
in  response  to  needs  for  electric  power  genera- 
tion. 

Region  3  (stations  15-19)  begins  just  below 
Region  2  and  continues,  also  for  more  than  100 
miles,  to  near  the  Massachusetts  border.  Within 
this  region  water  levels  are  also  controlled  for 
power  generation  and  lake-like  impoundments 
occur  in  a  nearly  unbroken  series.  With  the  ex- 
ception of  one  small  natural  area,  in  which  a 
native  mussel  fauna  of  at  least  5  species  still  sur- 
vives, the  fauna  throughout  the  whole  region 
has  apparently  been  reduced  to  one  species, 
Elliptio  complanata.  The  single  unspoiled  locali- 
ty below  Sumner's  Falls  in  Hartland,  Vt.  (sta- 
tion 17)  is  probably  the  site  of  the  several 
historical  records  from  Hartland  (see  Johnson, 
1915).  Here  the  river  tumbles  over  a  broad 
escarpment  and,  some  distance  below  the  falls 
where  reoxygenated  water  loses  its  turbulence, 
a  small  population  oi Alasmidonta  heterodon  ex- 
ists and  large  populations  of  other  species  still 
flourish.  Convenient  access  here  is  only  possible 
by  boat  and  then  only  during  the  infrequent  and 
unpredictable  periods  of  computer  permitted 
low-flow.  It  is  hoped  that  this  inaccessability  will 
contribute  to  the  survival  of  A.  heterodon  here 
and  to  the  conservation  of  the  unusually  healthy 
mussel  community  in  which  it  occurs. 

It  is  interesting  that  the  ubiiiuitous  and  abun- 
dant species  Elliptio  comphmato  is  entirely 
absent  from  Region  1,  a  situation  which  we  first 


Vol.  100(2) 


April  80,  1986 


THE  NAUTILUS      53 


ol)served  in  1955.  To  my  knowledge  there  are 
few  other  substantial  areas  within  the  g-eo- 
graphical  range  of  that  species  (northern 
Ontario  and  Nova  Scotia  to  Georgia),  where 
ecological  diversity  occurs  and  mussels  are 
found,  which  do  not  contain  E.  coniplanata.  It 
appears  probable  that  the  absence  of  E.  coni- 
planata  is  attributable  to  the  inability  of  its  only 
known  host  fish,  the  ye'low  perch,  to  ascend  the 
turbulent  and  shallow  portion  of  the  Connecti- 
cut River  below  the  Second  Connecticut  Lake. 
In  summary,  our  work  has  indicated  that  the 
upper  Connecticut  River  now  supports  mussel 
populations  of  good  species  diversity  in  only  two 
small  areas,  viz.  the  uppermost  Connecticut 
Lakes  and  their  vicinity,  and  an  unusual  locality 
at  Hartland,  Vt.  Other  work  in  1983,  which  is 
still  incomplete,  demonstrates  a  similarly  bleak 
but  uneven  situation  for  the  lower  Connecticut 
River.  The  diverse  mussel  community  at  South 
Hadley,  Massachusetts,  which  we  observed  in 
1952  is  now  gone  and  only  a  few  specimens  of 
Ellipflo  complanata  now  occur  there.  A  healthy 
molluscan  community  containing  E.  covtphntn- 
ta.  An.  rataracta,  A)h  iniplicafn.  and  AI.  U7i- 
ihiUitii  and  the  pleurocerid  GiDiinbusis  virginicd 


still  occurs  in  a  short  reach  of  the  Connecticut 
River  from  the  southern  l^order  of  Massachu- 
setts to  Warehouse  Point,  Connecticut, 
however. 

Clearly  the  mussel  fauna  of  the  whole  Connec- 
ticut River  now  survives  in  only  a  very  few 
remnant  communities.  Conservationists  and 
government  agencies  are  urged  to  help  in 
reversing  this  unfortunate  trend  toward 
extinction. 


LITERATURE  CITED 

Clarke,     Arthur    H..     1981.    The    Trilie    .Alasmidontini 

(Unionidae:  Anodontinae),  Part  1;  Piyuis,  Atasmidonta. 

and   Arridpris.    Smithsovinn    ('imtrihutions    to  Zoology, 

No.  326. 
Clench,  W.  J.  and  H.  1).  Russell.  1939.  Freshwater  Shells  of 

New   Hampshire.   Bioloyical  Surivy   of  the  Merrimuck 

Watfrsihed.  New  Hampshire  Fish  and  Game  Department, 

Surve.v  Report  No.  3,  pp.  201-206. 
1940.  Freshwater  Shells  of  New  Hampshire.  The 

NautUw!  54:52-53. 
Johnson,  Charles  W.,  1915.  P\-iuna  of  New  England.  List  of 

the  Mollusca.  Uccii!<i(iiiol  I'npfrx  of  the  Bustoti  Society  of 

Natural  History,  vol.  7,  part  13. 
Walker.  R.  D.  and  W.  H.  Cooli<lge.  Jr..  1908.  Mollusca  of 

Keene.  New  Hampshire.  The  NmitihiK  22:32. 


MORPHOLOGICAL  ANOMALIES  IN  THE  SHELL  OF 
FIELD-COLLECTED  BIOMPHALARIA  GLABRATA  (SAY,  1818) 


Jose  Gomez,  Mercedes  Vargas 

Departamento  de  Microbiologia  y  Parasitologia 

Universidad  Autonoma  de  Santo  Domingo 

Dominican  Republic 


and  Emile  A.  Malek 

Department  of  Tropical  Medicine 

Tulane  University  Medical  Center 

New  Orleans  70112-2699 


ABSTRACT 

We  report  the  occurrence  of  phenotypic  abnormal  shelU  of  the  freshwater  .swj.i/ 
Biomphalaria  glabrata  (Stt^,  1818)  in  different  habitats,  actual  and  potential  foci 
of  transmission  of  schistosomiasis  mansoni.  in  the  Dotninican  Republic.  There  is 
the  possibility  that  genetic  factors  cafuae  the  abnormalities  in  the  shell,  and  that 
they  are  inherited  through  cross-fertilization. 


Introduction 

During  the  normal  growth  of  a  shell  of  a  com- 
mon planorbid  snail  it  increases  in  diameter  in 
the  same  plane,  by  the  addition  of  new  whorls 


one  at  the  side  of  the  other  resulting  in  a  dis- 
coidal  form  (planispiral).  Sometimes,  however, 
this  growth  process  shows  anomalies  which  pre- 
sent the  malacologist  with  problems  in  the  iden- 


54      THE  NAUTILUS 


April  80,  1986 


Vnl.  100(2) 


TAl-iLK  I.  I 'ri'valencc  i)f  abiKirmai  shi'lls  aiiKing  natural 
populations  of  Biomfihiilnriti  gliiltralii  in  the  Dominican 
Republic. 


No. 

No, 

(%) 

No.  shells 

normal 

abnormal 

Locality 

examined 

shells 

shells 

Higuey 

124 

111 

13 

110  4) 

Sabana  de  la 

Mar 

209 

182 

27 

(13) 

Nisibon 

46 

46 

0 

Haina 

156 

153 

3 

(2) 

Jarabacoa 

48 

44 

4 

(8.3) 

Hato  Mayor 

226 

212 

14 

16) 

San  Francisco 

de 

Macoris 

25 

24 

1 

(41 

tification  of  a  particular  species.  Richards  (1970, 
1971)  observed  adult  BiomphaUirin  gluhmtn 
with  projecting  spires  in  genetic  stocks  showing 
frequent  embryos  with  the  first  whorl  open,  and 
with  high  eml)ryonic  mortality.  Phenotypic 
anomalies  in  the  shell  have  also  been  reported  by 
Andrade  and  Carvalho  (1973)  in  B.  straminea 
(Dunker,  1848)  raised  in  the  laboratory  in  Brazil 
and  by  Gomez  (1973)  in  B.  glabnUa  collected  in 
the  Dominican  Republic.  Andrade  and  Carvalho 
stated  that  deformation  of  the  last  whorl  was 
the  most  frequent  anomaly,  while  Gomez 
observed  abnormalities  of  various  types. 


In  this  paper  we  report  on  phenotypic  defor- 
mations in  the  shell  of  5.  glabraUx  in  populations 
occurring  in  various  actual  and  potential  foci  of 
transmission  of  schistosomiasis  mansoni  in  the 
Dominican  Republic.  A  total  of  834  shells  were 
examined  and  they  were  obtained  from  the  fol- 
lowing localities:  124  from  Higuey,  209  from 
Sabana  de  la  Mar,  46  from  Nisibon,  156  from 
Haina,  48  from  Jarabacoa,  226  from  Hato 
Mayor,  and  25  from  San  Francisco  de  Macoris. 

Results 

As  shown  in  Table  1,  of  124  snails  collected  in 
Higuey  13  demonstrated  conchological  anoma- 
lies for  a  prevalence  of  10%.  Among  209  snails 
which  came  from  Sabana  de  la  Mar,  27  (13%) 
were  abnormal,  whereas  those  from  Nisibon  did 
not  show  any  deviations.  Three  of  the  156  snails 
collected  in  Haina  exhibited  shell  abnormalities, 
as  well  as  4/48  from  Jarabacoa,  14/226  from 
Hato  Mayor,  and  1/25  from  San  Francisco  de 
Macoris. 

The  most  frequent  anomaly  was  the  deflection 
of  the  last  whorl  to  the  left  and  the  inclination  of 
the  preceding  whorl  to  the  right  (Fig.  1).  In 
some  shells  the  deflection  of  the  last  whorl  was 
pronounced  to  about  180°,  in  such  a  way  that 


4#  A 


mm 


FIG.  1.  liiniiiphdldnii  (iliihriitii  froni  the  Doniinican  Republic.  Top  row;  normal  shells,  left 
(under  or  umbilical)  side.  Middle  row;  abnormal  shells,  left  (under  or  umbilical)  side.  Bottom 
row;  abnormal  shells,  rinht  (upper)  side. 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      55 


the  aperture  was  detached  from  the  preceding 
whorl,  giving  the  shell  a  helicoidal  api)earance. 
As  a  result  the  early  whorls  overlapped,  and  the 
umbilicus  became  much  narrower  than  in  nor- 
mal shells.  In  addition  to  the  abnormalities  in 
the  form  of  the  shell  it  was  common  to  observe 
the  deposition  and  incrustation  of  calcareous 
material  in  the  abnormal  shells.  Scarifications  of 
the  umbilicus  were  never  observed. 

Discussion 

Sturrock  and  Sturrock  (1971)  showed  that  in 
St.  Lucian  B.  glabrata  infected  with  Schisto- 
soma mansoni  there  is  a  pronounced  distortion 
of  the  aperture  to  the  right  (upper)  side,  that  is, 
distortion  in  the  opposite  direction  from  that  of 
the  shells  we  are  reporting  upon  from  the 
Dominican  Republic.  The  distortion  in  the  in- 
fected snails  reported  by  the  above  authors, 
evidently  resulting  from  anatomical  changes 
due  to  the  infection  rather  than  being  genetical, 
first  appeared  at  the  end  of  the  prepatent  period 
of  the  infection  and  were  associated  with  the 
liberation  of  cercariae  from  the  snail. 

In  his  studies  on  the  genetics  of  B.  glahrata. 
Richards  (1970,  1973)  indicated  that  five  single 
gene  characters  have  been  demonstrated,  name- 
ly, basic  pigmentation,  pearl  formation,  antler 
tentacles,  everted  preputium  and  swollen  tenta- 
cles, and  adult  insusceptibility  to  infection  with 
S.  mansoni.  In  addition  to  the  above,  several 
multifactorial  inherited  growths  have  been 
observed  including  pulmonary  cavity  growths, 
tentacle  and  eye  variations  and  median  head 
bulbs.  Other  multifactorial  genetic  characters 
include    ajiertural    lamellae,    spire    formation 


(Richards,  1971),  mantle  pigment  and  suscepti- 
bility of  juvenile  B.  glabrata  to  infection  with  S. 
mamoni  (Richards  and  Merritt,  1972).  The 
presence  of  abnormal  shell  growth  in  different 
natural  populations  of  B.  glabrata  in  the 
Dominican  Republic  suggests  that  genetic  fac- 
tors cause  the  formation  of  these  anomalies.  The 
alisence  of  such  factors  in  certain  populations 
(Nisibon)  and  their  low  frequency  in  others  may 
indicate  their  association  with  a  sublethal  gene 
whose  penetration  may  inhibit  high  frequencies. 
However,  cross-breeding  experiments  are 
needed  to  determine  the  validity  of  this  specu- 
lation and  the  type  of  genetical  inheritance  in- 
volved in  the  formation  of  abnormal  shells. 

LITERATURE  CITED 

Andrade,  R.  M.  and  Carvalhu,  C.  S.  1973.  Aiiomalias  en 
conchas  de  Biamphaldrin  straminen  (Dunker.  1848), 
criadas  en  laboratorin  (Pulmonata.  Planorbidae).  Rev. 
SaudePuhlica  (S.  Pniilo)  7:28.3-284. 

Gomez.  .1.  197.3.  Distrihucion  geografica  del  Birmiphiilaria 
gliibrata  (huesped  intermediario  del  Sc/N.sfo.sowK  miDhsorii) 
en  Republica  Dominicana.  Tke^ix  for  Licenciado  en 
Biologia,  Universidad  Autononia  de  Santo  Domingo. 

Richards.  C.  S.  1970.  Genetics  of  a  molluscan  vector  of  schis- 
tosomiasis. Nature  227:806-810. 

1971.  Biiimphalnriri  (jUihnitu  genetics:  .Spire  for- 
mation as  a  sublethal  character.  Ji>iir.  Inrert.  Pnth. 
17:.53-i58. 

1973.  Genetics  of  Bior/ijihulnrin  glahmta  (Gas- 
tropoda: Planorbidae).  Mtilaeol.  Reriew  6:199-202. 

and  Merritt.  J.  W.  1972.  Genetic  factors  in  the 


susceptibility  of  juvenile  Bumiphalnrin  qlabrnta  to 
Schistosomii  in<ntxi)ni  infection.  Aiiiei'.  ■Imir.  Trap.  Mnl. 
Hyg.  21:425-434. 
Sturrock,  R.  F.  and  Sturrock,  B.  M.  1971.  Shell  abnormali- 
ties in  Biomphalnriii  gliibnila  infected  with  .Sf/(i.s(o.somfi 
//(((H.s-on/  and  their  significance  in  field  transmission 
studies,  .in// r.  Hehiiiiith<il.  45:201-210. 


NEW  MOLLUSK  JOURNALS 


Heldia,  Miinchner  Malakologische  Mitter- 
lungen,  a  West  German  mollusk  journal,  under 
the  editorship  of  Gerhard  Falkner  (ISSN 
0176-2621)  was  launched  in  1984.  Annual  sub- 
scription is  DM  25  (about  U.S.  $12.00).  Write: 
Postfach  26  01  23,  8000  Munich,  West  Germany. 
Most  articles  in  Band  1  have  been  on  land  and 
freshwater  moUusks,  but  some  new  marine 
species  are  described  {Conus  montillai  Rockel, 
1985). 

A-pex.  Informations  Scientifiques  de  la  Societe 


Beige  de  Malacologie  (a  quarterly),  vol.  1.  n<i.  1, 
36  pp.,  3  pis.  Editor:  R.  Duchamps.  Annual 
suliscription  of  900  B.F.  for  Apex  and  Arion 
(their  news  and  popular  publication)  should  be 
sent  by  international  money  order  or  bank  check 
in  Belgium  Francs  to:  M.  J.  Buyle,  Av.  M. 
Maeterlinck,  56,  bte  8,  Bruxelles,  Belgium 
B-1030.  The  first  numlier  contains  new  marine 
Rumanian  gastropods  by  Grossu  and  new 
Australian  X'olutidae  by  Foppe. 


56      THE  NAUTILUS 


April  80,  1986 


Vol.  100(2) 


THE  ECOLOGY  OF  FRESHWATER  GASTROPODS 
IN  THE  CENTRAL  CANADIAN  REGION 

Eva  Pip 

Department  of  Biology 

University  of  Winnipeg 

Winnipeg,  Manitoba 

Canada  R3B  2E9 

ABSTRACT 

The  ecology  of frefihwater  gastropods  was  surveyed  at  more  than  1,00  permnnent 
aquatic  habitats  located  within  the  area  bounded  by  47°  and  54°  N  and  H°  and 
106°  W  during  1972  to  1984.  Parameters  examined  were:  water  body  type,  sedi- 
ment type,  gastropod  community  diversity,  submerged  macrophyte  diversity,  sur- 
face water  pH,  total  dissolved  solids,  total  alkalinity,  chloride,  sulphate,  nitrate/ 
nitrite  and  soluble  reactive  phosphorus.  Many  species  showed  distributioTis  with 
respect  to  these  parameters  that  differed  from  the  total  site  sampling  distribu- 
tions. Gastropods  in  the  study  area  appeared  to  form  a  continuum  according  to 
habitat  type,  with  Stagnicola  palustris,  Physa  jennessi  and  Fossaria  modicella  at 
the  higher  extremes  of  water  chemistry,  and  S.  reflexa,  Bulimnea  megasoma, 
Marstonia  decepta  and  Helisoma  campanulatuni  at  the  lower  end  of  the  scale. 
Species  able  to  tolerate  high  dissolved  inorganic  values  also  showed  broad 
tolerance  ranges  for  these  parameters.  Regional  variation  in  tolerance  ranges 
may  be  a  com ple.r  function  of  adaptation  combined  with  the  inter-relationsihips 
between  various  aspects  if  the  habitats  themselves. 


The  physical  environment  is  believed  to  ac- 
count for  much  of  the  observed  distribution  pat- 
terns of  freshwater  gastropods  (e.g.  Aho,  1966). 
The  environmental  factors  governing  molluscan 
occurrence  have  been  the  sulyect  of  many 
studies  in  various  parts  of  the  world.  Of  par- 
ticular importance  have  been  the  contributions 
of  Macan  (1950,  1957)  and  Dussart  (1976)  in 
England,  Okland  (1979)  in  Norway  and  Aho  et 
al.  (1981)  in  Finland.  In  North  America  much 
ecol(jgical  intVirmation  has  been  presented  by 
Ilarman  and  Berg  (1971),  Clarke  (1973,  1981), 
Pip  (1978,  1985)  and  others. 

Various  aspects  of  the  aquatic  environment 
may  influence  snail  distributidii,  for  example 
water  chemistry,  degree  of  water  flow  or  tur- 
bulence, water  body  size,  substrate  type  and 
biotic  factors  such  as  type  and  abundance  of 
aquatic  vegetation  and  composition  of  the 
gastropod  community  (as  competition  may  be  a 
factor,  e.g.  Aho  et  al.  (1981)).  'i'lie  problem  is 
complicated  further  in  that  local  differences 
may  exist  in  the  tolerance  ranges  of  a  given 
species  with  respect  to  a  particular  parameter. 


and  thus  the  relative  importance  of  different 
parameters  may  show  regional  variation  (Pip, 
1985).  However,  such  variation  is  difficult  to 
detect  and  evaluate  since  few  data  are  available 
in  the  literature  in  consolidated  form  regarding 
the  ecological  characteristics  of  individual 
species  in  different  geographical  areas. 

The  present  paper  presents  the  results  of 
ecological  surveys  carried  out  from  1972  to  1984 
at  more  than  400  permanent  aciuatic  habitats 
located  within  the  area  bounded  by  47°  and 
54°N  and  94°  and  106°W  (eastern  Saskat- 
chewan, Manitoba,  northwestern  Ontario, 
northern  North  Dakota  and  Minnesota),  with 
sampling  locations  being  more  concentrated  in 
the  eastern  portion  of  this  area.  The  parameters 
examined  included  water  chemistry  (pH,  total 
dissolved  solids,  total  alkalinity,  chloride, 
sulphate,  nitrate  and  nitrite,  and  soluble  reac- 
tive phosphorus),  water  body  size  and  type, 
substrate  type,  and  diversity  and  species  com- 
position of  the  gastropod  and  submerged 
acjuatic  macrophyte  communities.  Analyses  of 
the  latter  parameter,  i.e.  community  composi- 


Vol.  100  (L 


April  80,  1986 


THE  NAUTILUS      57 


tion,  have  lieen  reported  elsewhere  (Pip,  1978, 
1985). 

Materials  and  Methods 

Surveys  were  conducted  during  May  to 
Sejitember  of  each  year.  All  sites  sampled  con- 
tained water  year-round  and  aquatic  communi- 
ties appeared  established.  Most  of  the  sites  were 
sampled  only  once.  However  11  sites  throughout 
the  region  were  monitored  for  water  chemistry 
parameters  at  2-week  intervals  during  an  entire 
sampling  season  in  order  to  estimate  the  range 
of  seasonal  variation  at  a  given  type  of  site.  Ap- 
proximately 50  additional  sites  were  scored  for 
all  parameters  at  least  twice  in  different  years. 

At  each  site  all  species  of  snails  encountered 
while  wading  or  canoeing  within  a  search  time 
of  1  hour  were  recorded.  Sul)merged  plants 
were  collected  by  hand  and  by  dredging  with  a 
rake;  these  were  taken  to  the  laboratory, 
washed  and  examined  for  additional  snails. 
Plant  community  diversity  was  scored  Ijy  in- 
cluding only  plants  that  were  at  least  partially 
sul)merged.  Large  water  bodies  were  sampled 
at  a  number  of  stations  and  each  was  treated  as 
a  separate  site  because  of  the  heterogeneity  of 
such  waters. 

A  surface  water  sample  was  collected  at  each 
site,  immediately  placed  on  ice  in  a  lightproof 
container  and  frozen  within  a  maximum  of  48 
hours  after  collection.  The  water  chemistry 
parameters  were  determined  on  the  thawed 
samples  using  methods  recommended  by  the 
American  Public  Health  Association  (1972).  The 
pH  was  measured  directly  in  the  field  using  a 
portable  pH  meter. 

When  compiling  the  water  chemistry  data,  if  a 
number  of  water  chemistry  values  were  avail- 
able for  the  same  site,  the  two  extreme  values  of 
each  parameter  at  that  site  were  included  in  the 
tolerance  range  of  the  species  which  occurred  at 
such  a  site. 

Results 

A  total  of  38  species  was  identified  as  occur- 
ring at  two  or  more  water  bodies  within  the 
study  area.  In  addition  to  these  identifiable 
species,  a  number  of  Physa  lots  were  collected 
whose  identity  could  not  be  established.  These 
were  of  several  morphological  types,  each  of 
which  was  present  at  a  number  of  sites. 


Water  body  tyjie  was  classified  according  to 
area,  if  lentic,  or  depth,  if  lotic  (Table  1).  The 
most  heterogeneous  group  was  that  of  ponds; 
this  group  included  water  bodies  of  various 
origins  and  basin  types,  such  as  natural  ponds, 
oxbows,  artificial  excavations  (ditches,  farm 
ponds,  (juarry  pits,  road  fill  excavations,  etc.), 
I)eaver  ponds,  spring  fed  pools  and  bog  pools. 

Many  gastropods  appeared  to  show  distribu- 
tions which  were  markedly  different  from  the 
overall  site  sampling  distribution.  In  all  subse- 
quent discussion,  only  species  for  which  N>5 
will  be  considered.  Although  almost  all  species 
could  occur  in  lakes,  this  water  body  type  was 
the  major  type  of  habitat  for  all  of  the  amni- 
colids,  and  ValiHita  spp.,  Stagnicola  catasco- 
pium,  Fossaria  decampi  and  Helisoma  carnpa- 
nulatum.  Ponds  were  the  major  water  body  type 
for  Stagnicola  caper ata,  Fossaria  exigua, 
Planorbula  campestris,  Armiger  crista,  and  to  a 
lesser  extent,  Stagnicola  palustris.  Aplexa 
hypnormn  and  Physa  jennessi  (ssp.  skinneri 
Taylor,  1953).  Bulim.nea  megasowM  appeared  to 
favor  rivers  above  other  habitat  types  when 
compared  to  the  overall  sampling  distribution. 
In  general,  running  waters  were  preferred  over 
lentic  habitats  by  Stagyiicola  rejlexa.  Bulimnea 
megasoma.  Ferrissia  rivularis  and  Planorbula 
armigera.  Gyraulus  deflectus  occurred  often  in 
both  running  waters  and  lakes  but  appeared  to 
avoid  ponds. 

Distribution  of  gastropods  with  respect  to 
substrate  type  (Table  2)  showed  that  Ferrissia 
rivularis  and  Stagnicola  reflexa  were  commonly 
found  in  granitic  Precambrian  Shield  basins, 
although  they  also  occurred  on  clay  and  coarse 
sediments  respectively.  Planorbula  campestris 
occurred  mostly  on  coarse  and  silt  sediments. 
Sand  appeared  to  lie  preferred  by  Valvata 
since7'a.  V.  tricarinata.,  Cincinnatia  cincinna- 
tiensis.  Marstonia  decepta.  Probythinella 
lacustris,  Stagnicola  catascopium,  Fossaria 
decampi,  Helisoma  campanulatum  and  Pro- 
nienetiis  umbilicatellus.  Fossaria  exigua  and 
Stagnicola  caperata  favored  both  sand  and  clay, 
while  clay  was  the  major  sediment  type  for 
Stagnicola.  palu.stris,  Fossaria  modicella  and 
Helisoma  pilsbryi.  Physa  jennessi  preferred 
both  silt  and  clay.  Physa  gyrina,  Lymnaea 
stagnalis.  Helisoma  anceps,  H.  trivolvis,  Planor- 
bula armigera  and  Gyraulus  parvus  appeared  to 


58      THE  NAUTILUS 


April  80,  1986 


Vol.  100(2) 


TABLE    I .   Percc'ntai;<'  ciislriliiiti<in  accunliiif;  t"  haliitat  tyin'  in  the  stiuly  area.  N  =  no.  ot'i)l)sena- 
tions. 


Species 


Lakes    Ponds    Rivers   Creeks     N 
>10  ha   <10  ha    -Zm  deep  '2ni  deep 


Campeloma  decisum  Say,  1816 
Valvata  sincere  Say,  1824 
V.  tricarinata  Say,  1817 
Cincinnatia  cincinnatiensi  s 

Anthony,  1840 
Marstonia  decepta  F.C.  Baker, 

1928 
Probythinella  lacustris  F.C. 

Baker,  1928 
Amnicola  1 imosa  Say,  1817 
A^  walkeri  Pilsbry,  1898 
Lymnaea  staqnal is  Linne,  1758 
Acella  haldemani  "Deshayes" 

Binney,  1867 
Bui imnea  megasoma  Say,  1824 


50 

50 

57 

29 

84 

14 

2 

73 

4 

23 

100 

86 

71 
80 
40 

100 
30 


Stagnicola  palustri  s  Muller,  1774  29 
S_.  catascopium  Say,  1817 
S^  caperata  Say,  1829 
S^  reflexa  Say,  1821 
Fossaria  da11i  F.C.  Baker,  1907 
1896 


92 

14 
29 
50 
75 
7 
35 
20 
47 
36 
22 


F.  decampi  Streng, 
F.  exigua  Lea ,  1841 

F.  modicel 1  a  Say,  1825 
F^   parva  Lea,  1841 
Physa  qyrina  Say,  1821 
P^  jennessi  Dall,  1919 
Aplexa  hypnorum  Linne,  1758 
Ferrissia  paral lela  Haldeman, 

1841 
F_^  rivularis  Say,  1817  45 

Hel isoma  trivolvis  Say,  1816  42 
H.  pi  1 sbryi  inf racarinatum 

F.C.  Baker,  1932  42 

FL  corpulentum  Say,  1824  50 
ti.  campanulatum  Say,  1821  81 
H^  anceps  Menke,  1830  47 

Planorbula  arniigera  Say,  1821  41 
P^  campestris  Dawson,  1875  29 
Promenetus  exacuous  Say,  1821  51 
P.  umbili  eaten  us  Cockerell,  1887  43 
Armiger  crista  Linne,  1758  33 
Gyraulus  parvus  Say,  1817  43 
G^  circumstriatus  Tryon,  1866     41 

G.  deflectus  Say,  1824  59 


14 


4 
7 

44 

26 


14 

7 

7 

19 

3 

69 

20 

5 

47 

6 

7 

246 
3 

35 

26 

9 

23 

57 

6 

8 

170 

8 

12 

86 

7 

29 

29 

13 

7 

25 

25 

4 

12.5 

12.5 

8 

86 

7 

14 

52 

9 

4 

46 

80 

5 

36 

10 

7 

253 

58 

6 

33 

62 

5 

11 

37 

100 

3 

19 

23 

13 

31 

43 

7 

8 

189 

37 

21 

19 

50 

4 

4 

15 

75 

44 

3 

6 

106 

29 

16 

14 

68 

71 

7 

31 

11 

7 

55 

43 

14 

7 

67 

9 

45 

5 

7 

161 

48 

4 

7 

27 

16 

14 

11 

44 

TOTAL  SITES 


40.5 


43.0 


9.0 


7.5   412 


be  substrate-indifferent. 

The  mean  gastropod  community  diversities 
where  each  species  was  found  were  higher  than 
the  mean  overall  site  diversity  (Table  3)  because 
all  species  except  Sfngnicola  rcjlexa  occurred  to 
some  extent  in  a  proportion  of  highly  diverse 


comnuinitii's.  For  species  with  N>5,  Fossaria 
ilcrnmiii  and  Frohnthinclla  hirustris.  both 
primarily  lacustrine  species,  showed  the  highest 
diversity  values.  Low  mean  (liversities  were 
seen  for  communities  in  which  Stagnicola 
rejlexa,  Ferrissia  rivularis.  Physa  gyrina.  Stag- 


Vol.  100(2)  April  30,  1986 

TABLE  2.   Perceiitaije  ilislriliutimi  acconiing  to  sul)strate  type  in  thf  study  area. 


THE  NAUTILUS      59 


Species 


Gravel/ 
Bedrock  coarse 

Granitic   Litnestone  Shale       sand 


Sand   Silt   Clay  Organic 


Campeloma  decisum 

75 

25 

4 

Valvata  sincera 

14 

57 

29 

7 

V.  tricarinata 

4 

2 

11 

46 

2 

20 

15 

46 

Cincinnatia  cincinnatiensi s 

15 

4 

62 

19 

26 

flarstonia  decepta 

88 

12 

8 

Probythinel la  lacustris 

29 

57 

14 

7 

Amnicola  limosa 

19 

12 

39 

1 

16 

13 

69 

A.  walkeri 

33.3 

33.3 

33.3 

6 

Lymnaea  stagnalis 

6.5 

2 

1 

17 

24 

6.5 

23 

20 

246 

Acella  haldemani 

100 

3 

Bulimnea  megasoma 

19 

19 

24 

14 

24 

21 

Staqnicola  palustris 

2 

2 

2 

21 

23 

9 

29 

12 

172 

S.  catascopium 

8 

8 

8 

50 

18 

8 

12 

S.  caperata 

57 

43 

7 

S.  reflexa 

43 

43 

14 

7 

Fossaria  dalli 

50 

25 

25 

4 

F.  decampi 

12.5 

75 

12.5 

8 

F.  exiqua 

7 

7 

43 

7 

36 

14 

F.  modicella 

2 

2 

2 

13 

22 

11 

34 

14 

45 

F.  parva 

20 

20 

20 

40 

5 

Physa  gyrina 

12 

2 

1 

15 

28 

5 

22 

15 

251 

P.  jennessi 

9 

21 

21 

34 

15 

33 

Aplexa  hypnorum 

3 

3 

34 

18 

5 

26 

11 

38 

Ferrissia  parallela 

33 

67 

3 

F.  rivularis 

37 

7 

13 

33 

10 

30 

Helisoma  trivolvis 

4 

2 

1 

18 

24 

9 

25 

17 

189 

H.  pilsbryi  inf racarinatum 

16 

4 

32 

32 

16 

19 

H.  corpulentum 

25 

25 

50 

4 

H.  catnpanulatum 

16 

1 

10 

49 

4 

6 

14 

73 

H.  anceps 

8 

19 

28 

2 

19 

24 

107 

Planorbula  armijera 

9 

3 

3 

15 

21 

3 

24 

22 

67 

P.  campestris 

43 

14 

29 

14 

7 

Promenetus  exacuous 

15 

4 

13 

16 

7 

29 

16 

55 

P.  umbilicatellus 

14 

14 

58 

14 

7 

Armiger  crista 

11 

23 

33 

33 

9 

Gyraulus  parvus 

5 

1 

1 

16 

26 

9 

21 

21 

161 

G.  ci rcumstriatus 

8 

4 

23 

34 

4 

8 

19 

26 

G.  deflectus 

12 

2 

9 

35 

5 

12 

26 

43 

TOTAL  SITES 

10.8 

2.0 

0.9 

15.9 

27.0 

5.9 

19.6 

17.9 

408 

nicoln  pdlusfris,  Lymnaea  stagnalis,  Helisoma 
anceps.  Bulimnea  m.egasnma.  Physa  jennessi. 
Fossaria  exigua  and  Helisoma  trivolvis  were 
present.  The  maximum  ret-onled  gastro|)od 
community  diversity  was  17. 

Mean  macrophyte  community  diversity  was 
high  for  Valvata  sincera.  Gyrauluji  deflectus. 
Hciisonut  campanulatum,  Bulimnea  megasonta 
and  all  amnicolids  except  Prohythine.lla 
lacuMris.  Low  mean  macrophyte  diversities  {<6) 
were  seen  for  Physa  jennessi,  Fossaria  modi- 
cella, Probythinella  laciistris.  Fossaria  exigua. 
Stagnicola  palustris  and  S.    capcrala.    Of  tlie 


above  species,  mean  diversity  values  were  low 
for  both  gastropod  and  macrophyte  communi- 
ties for  Physa,  jennessi.  Fossaria  exigua  and 
Stagnicola  i)alu.'<fris.  As  has  previously  been 
noted  by  Pip  (1985),  there  was  little  correlation 
between  the  two  kinds  of  diversity  at  the  dif- 
ferent site  types,  aside  from  the  tendency  of 
ponds  to  contain  few  species  of  both  snails  and 
plants. 

The  widest  range  for  surface  water  pH  was 
observed  for  Helisoma  trivolvis  (5.5  pH  units, 
equal  to  the  total  sampling  range)  (Table  4), 
followed    by    H.    fnireps.    Ferrissin    rivularis, 


60      THE  NAUTILUS 


Ai)ril  80,  1986 


Vol.  100(2) 


TABLE  3.   Moan  trastnip"<l  ami  suliiiiei'f;tMi  macniphyte  divorsily  of  ccimiminities  in  which  each 
species  was  found  in  the  study  area. 


Species 

Gastropod 
diversity 

N 

Macrophyte 
diversity 

N 

Campeloma  decisum 

13.3 

3 

17.3 

3 

Valvata  sincera 

9.6 

7 

10.4 

7 

V.  tiicarinata 

8.3 

46 

8.1 

46 

Cincinnatia  cincinnatiensis 

7.7 

27 

12.2 

25 

Marstonia  decepta 

7.3 

8 

14.6 

8 

Probythinella  lacustris 

10.0 

7 

5.0 

7 

Amnicola  limosa 

7.7 

68 

10.6 

68 

A.  walkeri 

6.0 

5 

11.6 

5 

Lymnaea  stagnalis 

5.5 

246 

6.9 

246 

Acella  haldemani 

11.0 

3 

16.7 

3 

Bulimnea  meqasoma 

5.6 

21 

10.1 

19 

Stagnicola  palustris 

5.5 

173 

5.4 

166 

S.  catascopium 

9.2 

13 

6.4 

11 

S.  caperata 

6.1 

7 

5.8 

6 

S.  reflexa 

4.0 

7 

9.9 

6 

Fossaria  dalli 

7.8 

4 

8.0 

4 

F.  decampi 

11.0 

8 

8.1 

8 

F.  exiqua 

5.8 

14 

5.4 

13 

F.  modicella 

7.1 

47 

5.0 

44 

F.  parva 

6.6 

5 

5.0 

5 

Physa  gyrina 

5.4 

250 

7.3 

246 

P.  jennessi 

5.7 

33 

4.8 

31 

Aplexa  hypnorum 

6.7 

36 

6.2 

33 

Ferrissia  parallela 

10.5 

2 

17.0 

2 

F.  rivularis 

5.3 

31 

7.6 

30 

Helisoma  trivolvis 

5.8 

189 

7.0 

189 

H.  pilsbryi  inf racarinatum 

7.2 

19 

9.0 

18 

H.  corjjulentum 

5.0 

4 

9.5 

4 

H.  campanulatum 

6.4 

74 

11.6 

73 

H.  anceps 

5.6 

108 

7.2 

102 

Planorbula  armigera 

6.7 

68 

8.5 

63 

P.  campestris 

7.7 

6 

3.7 

6 

Promenetus  exacuous 

7.7 

55 

7.6 

65 

P.  umbilicatellus 

4.9 

7 

8.9 

7 

Armiger  crista 

9.7 

9 

3.8 

9 

Gyraulus  parvus 

6.0 

165 

7.2 

158 

G.  circumstriatus 

6.6 

27 

7.5 

27 

G.  deflectus 

7.3 

44 

11.7 

41 

TOTAL  SITES 

4.6 

403 

6.8 

397 

Gyraulus  dejlertus,  Stagnix'olu  palusinx, 
Fossaria  rnodicella  and  Gyraulus  parvus  (all  at 
>  4  pH  units).  The  narrowest  ranj^es  (<  2  units) 
were  seen  for  Planorbula  campestris,  Mar- 
stonia decepta  and  Fossaria  decampi;  these 
three  species  liowever  were  sampled  at  relative- 
ly few  sites.  The  highest  sampled  pH  of  \()Ji  was 
tolerated  by  Stagnicola  palustris,  Fassaria 
modicella,  Helisoyna  trivohns,  Promenetus 
exacuous,  Aryniger  rj-ista  and  Gyraulus  parvus, 
while  the  lowest  uH  of  5.0  was  observed  for 


Bulimnea  megasonia.  Ferrissia  rivularis, 
Helisoma  trivolvis,  H.  anceps  and  Gyraulus 
dejlertus. 

The  widest  ranges  for  total  dissolved  solids, 
encompassing  nearly  the  total  range  sampled, 
were  seen  for  Fo.nsaria  modicella.  Stagnicola 
fiatustris  and  Physa  jennessi  (Table  4),  followed 
by  somewhat  smaller  ranges  (approx.  4500  mg/1) 
for  Lymnaea.  stagyialis  and  Helisoma  trivolvis. 
Narrow  ranges  (300  mg/1  or  less),  all  towards 
the   bottom   end  of  the  scale,   were  seen   for 


Vol.  100(2) 


April  :;o,  i;)S6 


THE  NAUTILUS      HI 


TABLK    1.    Minimuni.  iikixiiiiuiii  ami  luraii  \aliK'S  fcir  pH  ami  tutal  disscilvt'd  solids  for  ffaslnipods  in 
the  stud\  area. 


Species 


pH  Total 

dissolved  solids,  mg/1 
Min.   Max.    x    H  Min.   Max.    x    H 


Campeloma  dec i sum 

7.1 

8.1 

7.8 

3 

58 

123 

87 

3 

Valvata  sincera 

6.2 

9.2 

7.4 

6 

134 

384 

264 

6 

v.  tricarinata 

6.7 

9.8 

8.3 

48 

33 

1794 

432 

47 

Cincnnnatia  ci nci nnat lens  i  s 

7.3 

9.8 

8.3 

25 

33 

875 

170 

24 

Marstoma  decepta 

7.7 

9.2 

8.4 

9 

60 

208 

134 

7 

Probythinel la  lacustris 

7.9 

9.5 

8.5 

5 

104 

1300 

494 

5 

Amnicola  limosa 

6.1 

9.8 

8.1 

80 

16 

1794 

235 

79 

A.  wall<eri 

7.0 

9.0 

7.9 

6 

55 

219 

99 

5 

Lymnaea  staqnalis 

6.2 

10.0 

8.1 

260 

16 

4549 

337 

255 

Acella  haldemani 

7.4 

8.5 

8.0 

3 

68 

143 

111 

3 

Bulimnea  meqasoma 

5.0 

9.0 

7.2 

29 

16 

271 

93 

29 

Stagnicola  palustris 

6.2 

10.5 

8.2 

174 

38 

5533 

535 

170 

S.  catascopium 

7.3 

9.5 

8.6 

12 

37 

1300 

509 

11 

S.  caperata 

6.6 

8.4 

7.8 

3 

196 

919 

504 

3 

S.  reflexa 

6.6 

8.8 

7.7 

7 

16 

80 

58 

7 

Fossaria  dalli 

7.5 

8.3 

7.8 

3 

89 

280 

212 

3 

F.  decampi 

7.9 

9.5 

8.6 

6 

104 

1300 

354 

6 

F.  exiqua 

6.6 

9.8 

7.8 

14 

129 

1763 

371 

13 

F.  modi  eel  1  a 

6.2 

10.5 

8.3 

49 

33 

5533 

751 

47 

F.  parva 

6.7 

9.5 

8.1 

4 

152 

1596 

565 

4 

Physa  gyrnna 

6.1 

10.1 

8.0 

267 

16 

2399 

272 

261 

P.  jennessi 

7.1 

10.5 

8.4 

28 

60 

5533 

799 

28 

Aplexa  hypnorum 

6.6 

9.5 

7.8 

35 

37 

1402 

424 

35 

Ferrissia  parallela 

7.1 

7.3 

7.2 

2 

78 

219 

149 

2 

F.  rivularis 

5.0 

9.5 

7.6 

38 

15 

1300 

169 

37 

Helisoma  tnvolvis 

5.0 

10.5 

8.0 

197 

16 

4549 

345 

196 

H.  pilsbryi  i nf racari natum 

6.6 

9.5 

7.7 

15 

16 

743 

169 

15 

H.  corpulentum 

8.0 

8.4 

8.2 

5 

42 

80 

57 

5 

H.  campanulatum 

6.9 

9.5 

8.1 

91 

30 

334 

104 

89 

H.  anceps 

5.0 

9.8 

8.0 

113 

18 

1333 

237 

111 

Planorbula  armigera 

6.2 

9.7 

7.8 

68 

16 

1794 

339 

67 

P.  campestris 

7.5 

8.8 

7.9 

6 

138 

459 

273 

6 

Promenetus  exacuous 

6.9 

10.5 

8.2 

60 

40 

1794 

413 

60 

P.  umbi 1 icatel lus 

6.7 

9.4 

8.1 

6 

80 

734 

267 

6 

Armiger  crista 

7.3 

10.5 

8.4 

10 

219 

1794 

836 

10 

Gyraulus  parvus 

6.2 

10.5 

8.1 

176 

24 

2399 

317 

172 

G.  CT rcumstriatus 

6.6 

9.8 

8.3 

26 

18 

2108 

418 

26 

G.  deflectus 

5.0 

9.4 

7.8 

48 

32 

1571 

176 

47 

TOTAL  SITES 

5.0 

10.5 

8.0 

446 

10 

5533 

302 

440 

Stagnicola  reflexa,  Marstonia  decepta.  Valonta 
fiincera,  Bulimnea  m.egaso»ia  and  Helisoma 
campanulatum. 

Total  alkalinity  ranges  (Table  5)  were  widest 
(>  700mg/l  CaCOj)  for  Stagnicola  palustris. 
Gyraulus  deflectus,  G.  circumstriatus  and 
Physa  jennessi.  Narrow  ranges  (<100  nig/l 
CaCOs),  all  again  towards  the  bottom  end  of  the 
scale,  were  seen  for  Stagyiicola  reflexa  and 
Valvata  sincera.  both  of  which  were  repre- 
sented by  small  numbers  of  samples.  Thirteen 
species  occurred  at  alkalinity  values  as  low  as 


0-4  mg/1  CaCOa,  while  only  three  were  observed 
at  the  maximum  sampled  value  of  800  mg/1 
CaCOs. 

All  species  tolerated  chloride  values  of  0  mg/1 
(Table  5),  but  only  Stagnicola  palustris, 
Fossaria  modicella  and  Physa  jennessi  could  be 
found  throughout  the  complete  range  sampled. 
Valvata  sincera  and  Stagnicola  reflexa  ap- 
parently avoided  chloride  altogether,  although 
again  these  were  represented  by  small  sample 
numljers.  Fossaria  decampi  and  Marstonia 
decepta  showed  chloride  ranges  of  <  10  mg/1. 


62      THE  NAl'TILUS 


April  30,  1986 


Vol.  100(2) 


TABLE  5.   Mininiuiii.  niaxinium  and  mi'aii  valufs  tor  total  alkalinity  and  chloridi-  for  tjastrupods  in 
the  study  area. 


Species 

Total 

alkal 

i  ni  ty . 

,  mg/1 

) 

Chloric 

le,  mg/1 

CaCO., 

Min. 

Max. 

"  X 

N 

Min 

.  Max. 

x 

N 

Campeloma  decisum 

64 

128 

97 

3 

0 

0 

0 

3 

Valvata  sincera 

70 

166 

119 

6 

0 

0 

0 

6 

V.  tricarinata 

0 

448 

144 

47 

0 

480 

70 

47 

Cinci'nnatia  cincinnatiensis 

14 

239 

91 

24 

0 

18 

3 

23 

Marstonia  decepta 

70 

220 

122 

8 

0 

8 

1 

8 

Probythinel  la  lacustris 

70 

159 

132 

5 

0 

238 

48 

5 

Amnicola  limosa 

4 

448 

96 

78 

0 

480 

34 

77 

A.  walkeri 

11 

172 

63 

5 

0 

8 

2 

5 

Lymnaea  stagnalis 

0 

560 

135 

255 

0 

592 

25 

254 

Acella  haldemani 

72 

100 

81 

3 

0 

0 

0 

3 

Bulimnea  megasoma 

9 

172 

54 

29 

0 

56 

3 

28 

Stagnicola  palustris 

4 

800 

180 

172 

0 

1234 

51 

172 

S.  catascopium 

84 

184 

134 

11 

0 

430 

130 

11 

S.  caperata 

106 

224 

173 

3 

0 

7 

2 

3 

S.  reflexa 

0 

72 

31 

7 

0 

0 

0 

7 

Fossaria  dalli 

72 

202 

129 

3 

0 

4 

1 

3 

F.  decamgi 

70 

284 

138 

6 

0 

2 

0. 

3  6 

F.  exigua 

36 

560 

183 

13 

0 

77 

10 

13 

F.  modicella 

48 

448 

194 

48 

0 

1234 

115 

48 

F.  parva 

108 

230 

167 

4 

0 

21 

5 

4 

Physa  gyrina 

0 

560 

122 

263 

0 

592 

24 

262 

P.  jennessi 

62 

768 

196 

28 

0 

1234 

79 

28 

Aplexa  hypnorum 

4 

448 

179 

33 

0 

438 

17 

33 

Ferrissi'a  parallela 

128 

172 

150 

2 

0 

0 

0 

2 

F.  rivularis 

4 

220 

68 

38 

0 

238 

9 

37 

Hell  soma  trivolvis 

0 

654 

144 

195 

0 

480 

22 

195 

H.  pilsbryi  inf racarinatum 

10 

316 

106 

15 

0 

261 

18 

15 

H.  corpulentum 

10 

41 

29 

5 

0 

0 

0 

5 

H.  campanul  atum 

10 

280 

71 

89 

0 

15 

0. 

5  88 

H.  anceps 

4 

448 

115 

111 

0 

470 

23 

111 

Planorbula  armigera 

0 

560 

147 

67 

0 

480 

49 

67 

P.  campestris 

148 

280 

210 

6 

0 

13 

3 

6 

Promenetus  exacuous 

4 

654 

153 

60 

0 

480 

45 

59 

P.  umbilicatellus 

12 

266 

107 

6 

0 

22 

4 

6 

Armiger  crista 

80 

448 

241 

10 

0 

480 

146 

10 

Gyraulus  parvus 

0 

654 

134 

174 

0 

592 

28 

174 

G.  circumstriatus 

25 

800 

222 

26 

0 

602 

46 

26 

G.  deflectus 

20 

800 

99 

47 

0 

394 

13 

46 

TOTAL  SITES 

0 

800 

122 

440 

0 

1234 

24.4 

440 

All  species  (except  Stagnicola  caperata)  were 
observed  in  the  absence  of  measurable  sulphate 
(Table  6),  but,  as  for  chloride,  only  Sttignicota 
paliiHtris,  Fossaria  modicella  and  Physa.  jen- 
nessi were  found  throughout  the  entire  range 
sampled.  Valvata.  sincera.  Stagnicola  reflexa, 
Fossaria  decampi  and  Bulimnea  megasoma  ap- 
peared to  be  relatively  intolerant  of  sulphate. 

For  nitrate  and  nitrite  (Table  6),  Cincinnatia 
cincinnatiensis,  Planorbula  armigera  and  Stag- 
nicola palustris  occurred  throughout  all  or  most 
of  the  range  sampled.  Narrow  ranges  towards 


the    lower    end    of   the    scale    were    seen    for 
Stagnicola  re.jlexa.  and  Marstonia  decepta. 

Eleven  species  were  observed  to  tolerate  the 
highest  phosphorus  levels  (Table  7).  Bulimnea 
megasoma  and  Stagnicola  rcjlexa  appeared  to 
favor  low  phosphorus  concentrations. 


li 


Discussion 

the    study    area    only    niarginallv    more 


species  were  found  in  lakes  than  in  ponds. 
Although  none  of  the  species  appeared  to  be 
restricted    to    either    lakes    or    ponds,    many 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      63 


TABLE  ().   Miiiinuiiii,  niaxiiiiuni  .uid  mean  values  fur  sulphate  and  nitrate/nilrile  for  t;astrn|ioils  in 
the  study  area. 


Species 


Sulphate,  mg/l 
Min.   Max.    x 


Mi trate/nitri te,  mg/l 
N  Min.   Max.    x    N 


Campeloma  decisum 

0 

0 

0 

3 

0 

0.7 

3 

Valvata  sincera 

0 

2 

0. 

5  7 

1.0 

3.3 

1.6 

6 

V.  tricarinata 

0 

170 

51 

54 

0 

2.4 

1.3 

48 

Cincinnatia  c i nci nnat lensis 

0 

170 

21 

25 

0 

7.5 

1.1 

25 

Marstonia  decepta 

0 

109 

13 

9 

0 

1.5 

0.8 

9 

Probythinel la  lacustris 

0 

170 

69 

6 

0 

1.2 

0.5 

5 

Amnicola  limosa 

0 

170 

23 

85 

0 

2.7 

0.9 

78 

A.  walkeri 

0 

15 

4 

5 

0 

1.5 

0.9 

5 

Lymnaea  staqnalis 

0 

334 

28 

263 

0 

5.5 

1.2 

258 

Acella  haldemani 

0 

0 

0 

3 

0.8 

1.0 

0.9 

3 

Bulimnea  megasoma 

0 

15 

3 

30 

0 

5.3 

0.7 

28 

Stagnicola  palustns 

0 

3403 

65 

182 

0 

7.2 

1.6 

174 

S.  catascopium 

0 

113 

32 

12 

0.1 

1.7 

1.1 

12 

S.  caperata 

1.6 

157 

61 

3 

1.0 

2.3 

1.5 

3 

S.  reflexa 

0 

10 

2 

7 

0 

1.4 

0.5 

7 

Fossaria  dalli 

0 

39 

13 

3 

0.8 

1.3 

1.1 

3 

F.  decampi 

0 

14 

4 

6 

0 

2.4 

1.2 

6 

F.  exigua 

0 

161 

29 

14 

0.2 

3.3 

1.0 

14 

F.  modicella 

0 

3403 

129 

49 

0 

5.6 

1.6 

49 

F.  parva 

0 

148 

37 

4 

0.2 

2.4 

1.1 

4 

Physa  qyrina 

0 

334 

24 

271 

0 

5.5 

1.1 

265 

P.  jennessi 

0 

3403 

190 

28 

0 

6.8 

2.2 

28 

Aplexa  hypnorum 

0 

165 

32 

37 

0 

2.4 

1.3 

35 

Ferrissia  paral lela 

0 

104 

42 

3 

0.9 

1.5 

1.2 

2 

F.  ri  vulari  s 

0 

161 

18 

36 

0 

5.3 

0.6 

36 

Helisoma  trivolvis 

0 

443 

38 

204 

0 

5.2 

1.3 

197 

H.  pilsbryi  i nf racari natum 

0 

64 

5 

15 

0 

2.8 

0.7 

15 

H.  corpulentum 

0 

5 

2 

5 

0.1 

1.4 

0.6 

5 

H.  campanul atum 

0 

108 

3 

92 

0 

2.4 

0.8 

90 

H.  anceps 

0 

167 

18 

114 

0 

5.3 

1.1 

112 

Planorbula  armiqera 

0 

157 

25 

70 

0 

7.5 

1.3 

68 

P.  campestns 

0 

115 

45 

6 

0.9 

1.6 

1.2 

6 

Promenetus  exaciious 

0 

158 

42 

64 

0 

5.2 

1.3 

59 

P.  umbi 1 icatel 1  us 

0 

159 

28 

6 

0.3 

2.0 

1.2 

6 

Armiger  crista 

0 

128 

63 

11 

0.1 

2.9 

1.4 

10 

Gyraulus  parvus 

0 

170 

32 

181 

0 

5.2 

1.2 

176 

G.  CT rcumstri atus 

0 

158 

35 

26 

0.7 

6.8 

2.0 

26 

G.  deflectus 

0 

108 

8 

49 

0 

5.3 

1.2 

47 

TOTAL  SITES 

0 

3403 

31 

456 

0 

7.5 

1.1 

445 

gastropods  showed  strong  tendencies  to  occui- 
in  one  of  tiiese  liahitat  types.  Aho  rt  al.  (19<S1) 
oliserved  a  similar  situation  in  Finland,  where 
most  snails  could  be  found  over  a  wide  range  of 
water  body  sizes.  The  majority  of  the  species  en- 
countered in  the  present  study  were  quite 
plastic  in  terms  of  the  water  body  and  substrate 
type  in  which  they  could  occur.  Such  plasticity 
would  be  exjiected  to  be  advantageous  in  that 
the  species'  potential  for  colonization  of  new 
haliitats  would  be  enhanced. 

Most  of  the  species  (excejit  perhaps  .S/ai/i/zco/f/ 
reflexa)  could  function  in  highly  diverse  gastro- 


pod cnnimunities,  but  it  is  not  surprising  that 
many  species  were  found  frequently  in  less 
diverse  groupings  where  interspecific  competi- 
tion would  be  reduced.  Species  found  often  at 
low  community  diversities  were  capalile  of  sur- 
viving in  more  extreme  habitats  where  other 
species  failed.  Species  with  low  mean  communi- 
ty diversity  values,  such  as  Stagnirold  rejle.nt 
and  BiiVuitnea  tnefid.^oma.  could  occur  in  ex- 
treme habitats  only  at  the  lower  end  of  the  scale 
for  most  chemical  parameters  and  their  toler- 
ance ranges  for  these  parameters  were  narrow. 
On  the  other  hand,  species  stich  as  Stniinicula 


64      THE  NAUTILUS 


April  30,  1986 


Vol.  100  (2) 


TABLE  7.  Miniiiuim.  maximum  and  moan  values  tor 
molybdenum  reactive  phosphorus  for  gastropods  in  the 
study  area. 


Species 

Molybdenum  react 

.ive  P 

,  mq/l 

Min. 

Max. 

X 

N 

Campeloma  decisum 

0.48 

1.6 

0.9 

3 

Valvata  sincera 

0.23 

7.0 

2.3 

6 

V.  tricarinata 

0 

44.0 

6.1 

47 

Cinclnnatia  cinci nnatiensis 

0 

14.9 

2.9 

23 

Marstonia  decepta 

0 

5.6 

3.2 

9 

Probythinel la  lacustris 

1.7 

15.0 

7.1 

5 

Amnicola  limosa 

0 

44.0 

4.1 

78 

A.  walkeri 

0 

3.6 

1.6 

6 

Lymnaea  staqnalis 

0 

44.0 

3.4 

258 

Acella  haldemam 

0.60 

6.3 

3.5 

3 

Bulimnea  meqasoma 

0 

2.3 

0.7 

27 

Stagnlcola  palustris 

0 

44.0 

4.4 

174 

S.  catascopium 

0.45 

15.0 

3.3 

12 

S.  caperata 

0.51 

3.2 

2.3 

3 

S.  reflexa 

0 

2.4 

0.7 

7 

Fossaria  daUi 

0.74 

5.0 

2.3 

3 

F.  decampi 

0.56 

15.0 

4.7 

6 

F.  exiqua 

0.04 

5.5 

1.3 

14 

F.  modicella 

0 

44.0 

6.5 

49 

F.  parva 

0 

3.1 

1.1 

4 

Physa  qyrina 

0 

44.0 

3.4 

265 

P.  jennessi 

0 

15.7 

3.4 

28 

Aplexa  hypnorum 

0.04 

21.3 

6.1 

35 

Ferrissia  parallela 

0.62 

1.6 

1.  1 

2 

F.  rivularis 

0 

15.7 

2.4 

35 

Hel  Tsoma  tn  vol  vis 

0 

44.0 

3.6 

197 

H.  pilsbryi  inf racarinatum 

0.04 

5.6 

1.1 

15 

H.  corpulentum 

0.46 

1.8 

0.9 

5 

H.  campanulatum 

0 

10.6 

2.2 

90 

H.  anceps 

0 

21.3 

2.5 

112 

Planorbula  armiqera 

0.21 

44.0 

5.1 

68 

P.  campestris 

4.0 

12.4 

7.0 

6 

Promenetus  exacuous 

0 

44.0 

5.1 

59 

P.  umbi 1 icatel lus 

0.35 

8.3 

3.9 

6 

Armiqer  crista 

0.62 

44.0 

15.7 

10 

Gyraulus  parvus 

0 

44.0 

3.6 

176 

G.  circumstriatus 

0.14 

12.4 

3.0 

26 

G.  deflectus 

0 

10.6 

2.6 

46 

TOTAL  SITES 

0 

44.0 

2.2 

445 

paluHtrin,  Fhysd  jniHf'ss),  Hciisiinid  fricoh'is,  IL 
anceps,  Lyrrmaea  stagnnlh  and  Fen-tssln 
nvularis  could  occur  at  very  wide  ranges  of 
some  or  all  of  the  parameters  examined.  Some 
of  the  species  in  the  latter  s^rouji  were  also  very 
common  in  the  study  area,  suj^gestinj^  that,  in 
conjunction  with  their  ability  to  tolerate  a  wide 
range  of  conditions,  they  may  also  l)e  effective 
colonizers  of  new  habitats. 

A  low  value  for  macrophyte  community  divei'- 
sity  may  be  indicative  of  extreme  habitat  condi- 
tions and/or  of  new  habitats  in  which  succession 
has  not  peaked.  BuHnnwd  incfidsonui.  which 
showed  a  low  gastropod  and  high  macrophyte 
community  diversity,  was  characteristic  of 
habitats  in  the  eastern  portion  of  the  study  area, 
where  water  parameter  values  may  be  extreme- 
ly low  but  plant  diversities  are  high  because  of 
the  large  nutiibers  of  macrophyte  species  which 


occur  in  the  Precambrian  Shield  region  (Pip, 
1979,  1984).  Low  diversity  values  for  both 
gastropods  and  macrophytes,  as  seen  for  Phym 
jennessi.  Fossaria  exigua  and  Stagnicola 
palustris.  probably  reflected  not  only  extreme 
habitat  conditions,  but  also  the  pond  water  body 
ty])e  which  was  predominant  for  these  species. 
Macrophyte  diversity  is  characteristically  low  in 
ponds.  Some  regional  differences  in  both  gastro- 
pod and  macrophyte  community  diversities  ap- 
pear to  exist  for  a  given  snail  species  (Pip,  1985). 

The  species  present  in  the  study  area  formed  a 
continuum  with  respect  to  the  habitat  types  in 
which  they  were  found.  At  one  end  of  this  con- 
tinuum could  be  placed  Stagnicola  palustris, 
Physa  jennessi  and  Fossaria  modicella.  These 
species  frequented  ponds  and  a  clay  substrate. 
The  variability  of  water  quality  in  such  habitats 
was  associated  with  very  wide  tolerance  ranges 
of  these  species  for  pH,  total  dissolved  solids, 
total  alkalinity,  chloride  and  sulphate. 

At  the  other  end  of  the  scale  were  Stagnicola 
rejlexa,  Bulimnea  ynegasorna,  and,  to  a  lesser  ex- 
tent, Marstonia  decepta.  The  first  two  species 
were  found  most  frequently  in  lotic  habitats, 
while  the  last  species  was  found  in  lakes.  All 
were  found  primarily  in  Precambrian  Shield 
habitats  and  were  characterized  by  narrow 
tolerance  ranges  at  the  lower  end  of  the  scale 
for  total  dissolved  solids,  total  alkalinity, 
chloride,  sulphate,  nitrate  and  nitrite  and 
phosphorus.  Mean  parameter  values  were  also 
low  for  Helisoma  campanulatum,  although  this 
species  could  tolerate  somewhat  broader  ranges 
for  some  parameters  than  could  the  latter  three 
species,  and  possible  because  of  this  reason  it 
was  the  commonest  of  this  group.  These  species 
may  be  regarded  as  oligotrophic/dystrophic 
specialists. 

In  general,  species  which  could  tolerate  high 
water  chemistry  parameter  values  tended  to 
have  wide  tolerance  ranges.  Although  species 
with  wide  tolerance  ranges  coulcl  survive  in 
hal)itats  with  lower  parameter  values,  they  fre- 
(luenled  sites  with  higher  levels  of  dissolved  in- 
organics, as  seen  from  tlu'ir  mean  values. 
Species  such  as  .Arniiger  crista.  Physa  jcinn'ssi 
and  Fossaria  modicella  show  high  mean  values 
for  total  dissolved  solids  and  this  may  be  one 
reason  why  these  species  are  extremely  rare 
east  of  the  Precaml)rian  Shield  boundary,  where 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      65 


water  parameter  values  are  low.  Plnn(irl}itla 
campestris  may  also  belong  to  this  group  but 
was  insufficiently  sampled  in  the  present  study. 

A  number  of  workers  (e.g.  Aho,  1966,  1978; 
Dussart,  1976;  Okland,  1979;  Pip,  1978;  Dillon 
and  Benfield,  1982)  have  found  that  calcium 
hardness,  total  alkalinity  and  pH  are  important 
factors  in  determining  the  distribution  patterns 
of  many  freshwater  gastropods.  Okland  and 
Okland  (1980)  found  that  few  gastropods  occur- 
red below  pH  6.0  in  Norway,  with  only  a  few 
species,  not  stated,  being  able  to  tolerate  values 
as  low  as  5.2  (Okland,  1979).  Aho  (1978)  ob- 
served that  fewer  species  occurred  at  lower  than 
higher  pH  values  in  Finland.  The  findings  of  the 
present  study  supported  this  observation  up  to 
moderately  alkaline  values.  However  in  the 
study  area  five  species  could  occur  at  pH  values 
as  low  as  5.0.  Mackie  and  Flippance  (1983)  found 
that  in  southern  Ontario  Heli><oma  anceps  and 
Amnicola  limosa  could  occur  at  pH  5.5;  this 
record  for  the  latter  species  was  lower  than  the 
minimum  of  6.1  seen  in  the  present  study.  The 
latter  workers  also  reported  a  minimum  of  6.0 
for  Cincinnatia  cincinnatiensis  and  \'nlvata 
tricarinata.  compared  to  7.3  and  6.7  respective- 
ly in  the  present  study.  Ranges  for  all  other 
gastropods  reported  by  these  workers  fell 
within  the  ranges  obtained  in  the  present  study. 

Mackie  and  Flippance  (1983)  also  found  a 
lower  total  alkalinity  level  of  5  mg/l  CaCOj  for 
C.  cincinn.atieniiis  compared  to  a  value  of  14  in 
the  present  study.  All  other  ranges  found  by  the 
latter  workers  were  within  those  given  here. 

The  tolerance  range  of  a  given  gastropod 
species  for  a  particular  parameter  must  be 
viewed  in  the  context  of  the  geographical  area 
where  it  occurs.  Aho  (1966,  1978)  found  that 
with  respect  to  pH  and  calcium  content,  snail 
species  in  Finland  appeared  to  occur  in  lakes 
with  lower  values  for  these  parameters  than  in 
Sweden  or  Britain.  Similarly  Pip  (1985) 
reported  that  populations  in  Precambrian  Shield 
waters  often  seemed  to  tolerate  lower  values  of 
dissolved  inorganics  than  did  populations  to  the 
west  of  the  Shield  boundary.  Apparently  many 
species  may  undergo  adaptation  according  to 
the  water  cjuality  characteristics  predominant  in 
the  region. 

The  gastropod  communities  found  in  Precam- 
brian Shield  waters  in  the  study  area  appear  to 
tolerate  lower  minimum  pH   values  than   the 


communities  reported  by  Okland  (1979)  in 
Norwegian  lakes.  Perhaps  this  difference  is 
related  to  the  type  and  duration  of  acid  condi- 
tions in  the  two  areas.  The  acidity  of  undis- 
turbed Precambrian  Shield  lakes  is  derived 
largely  from  dissolved  organic  matter;  sulphates 
are  typically  absent.  The  acidity  of  the  lakes 
studied  by  Okland  (1979)  is  a  comparatively  re- 
cent development  as  a  result  of  acid  precipita- 
tion. Thus  snails  in  the  former  case  have  had 
more  opportunity  to  adapt  and  the  acidity  is  of 
different  chemical  origin.  Okland  (1979)  sug- 
gested that  the  importance  of  a  given  water 
ciuality  parameter  such  as  pH  may  be  related  to 
the  level  of  other  factors  present.  For  example 
Pip  (1985)  found  that  the  relative  importance  of 
specific  factors  may  change  in  different  geo- 
graphical regions.  Thus  the  tolerance  range  for 
a  particular  parameter  may  be  a  complex  func- 
tion of  genetic  and  physiological  traits  combined 
with  the  relationships  between  different 
chemical  entities  in  the  water  itself.  It  is  also 
possible  that  additional  habitat  characteristics, 
such  as  substrate  and  the  kinds  of  other 
organisms  present  in  the  community,  for  exam- 
ple vegetation,  may  further  mitigate  or  ag- 
gravate the  response  of  a  given  snail  population 
to  a  specific  parameter.  A  better  understanding 
of  the  factors  which  affect  snail  distribution  in  a 
given  area  can  only  be  achieved  when  compar- 
able data  becomes  available  for  a  number  of 
geographical  regions. 

LITERATURE  CITED 

Alio,  .1.  1966.  Ecological  liasis  of  the  distribution  of  the 
littoral  freshwater  molluscs  in  the  vicinity  of  Tampere, 
South  Finland.  Arm.  Zool.  Ferin.  3:287-322. 

1978.  Regional  variation  in  the  diversity  of  fre.sh- 


water  gastropods  in  southern  an<i  western  Finland.  Puhl. 
Utiif.  Joen.'iiiu.  Ser.  B,  No.  8,  10  pp. 

Aho.  .J.,  E.  Ranta  and  J.  Vuorinen.  1981.  Species  composi- 
tion of  freshwater  snail  communities  in  lakes  of  southern 
and  western  Finland.  ylwH.  Znnt.  Ferin.  18:233-241. 

Clarke,  A.  H.,  .Jr.  1973.  The  freshwater  molluscs  of  the 
Canadian  Interior  Basin.  Mnlacologia  13:1-509. 

1981.    The  Freshwater  MoUusrs   of  Cnnndii. 

National  Museums  of  Canada,  Ottawa.  446  pp. 

Dillon,  R.  T.  and  E.  F.  Benfield.  1982.  Distribution  of  pul- 
nionate  snails  in  the  New  River  of  Virginia  and  North 
Carolina,  U.S.A.:  interaction  between  alkalinity  and 
stream  drainage  area.  Fre^hunter  Biol.  12:179-186. 

Dussart,  G.  B.  J.  1976.  The  ecologj'  of  freshwater  molluscs 
in  north  west  England  in  relation  to  water  chemistry. 
J,,iir.  moll.  Sli«li,:'<  42:181-198. 

Harman,  W.  N.  and  C.  0.  Berg.  1971.  The  freshwater  snails 


66      THE  NAUTILUS 


April  :W.  1986 


Vol.  100(2) 


of  central  New  York  wiUi  illusli-atfd  keys  tci  the  n*'ii''''<i 
and  species.  Si'inrh:  Curnell  I'niv.  Ani'ic.  Kx|i.  Station 
1:1 -OS. 

Macan,  T.  T.  1  ;».')(>.  F^colnKX  <>t'  freshwater  Molhisca  in  the 
Engiish  Lake  District.  ./..»/•.  Anini.  EciA.  19:124-146. 

19.')7.  Chemical  analysis  in  ecokigj'  illustrateii 

front  Lake  District  tarns  ami  lakes.  3.  Faunistic  differ- 
ences. Priir.  Lnm.  Sm-  Lniiilmi  167: 172-I7"i. 

Mackie,  G.  L.  and  L.  A.  Khppance.  1983.  Intra-  and  inter- 
specific variations  in  calcium  content  of  freshwater  mol- 
lusca  in  relation  to  calcium  content  of  the  water.  Jitiir. 
imill.  &7h(//>.s' 49:204-212. 

Okland.  J.  1979.  Distriliution  of  environmental  factors  and 
fresh-water  snails  ((lastropoda)  in  Norway:  use  of  Rum- 
pean  invertebrate  survey  principles.  Muliu-olnijii:  18: 
211-222. 


Okland,  .1.  and  K.  A.  Okland.  19,S(l.  pH  level  and  food 
organisms  for  fish  ■  studies  of  UKK)  lakes  in  Norway.  In 
Proc.  Internat.  Conference  on  the  ecological  im|:iact  of 
acid  precipitation.  Norway.  1980  (D.  Drablos  and  \. 
Tollan,  eds.).  SNSF  Project,  pp.  326-327. 

Pip,  E.  1978.  A  survey  of  the  ecology  and  composition  of 
submerged  aquatic  snail-plant  ciminiunities.  Cniuididn 
■Inn,-.  Zoi.l.  56:22(;.3-2279. 

1979.    Survey    of   tlie    ecology    of   submerged 

aquatic  macro|ihytes  in  central  Canada.  AijikiHc  Hatdiiy 
7:339-357. 

1984.  Ecogeographical  tolerance  range  variation 


in  ai|uatic  macrophytes.  Hydrobidloyid  108:37-48. 

1985.  The  ecology  of  freshwater  gastropods  on 

the    southwestern    edge    of    the    Precambrian    Shield. 
Cariddinri  Fii'hl-Nitt.  99:76-85. 


PSEUDOSUCCINEA  COLUMELLA  (LYMNAEIDAE)  IN  THE  DOMINICAN 

REPUBLIC  AND  TRANSMISSION  OF 

FASCIOLIASIS  IN  THE  CARIBBEAN  REGION 


Jose  Gomez,  Mercedes  Vargas 

Departanieiitd  dv  Micrdhioiogia  y  Parasitologia 

Universidad  Autonoma  de  Santo  Domingo 

Dominican  Reiuihlic 


and  Emile  A.  Malek 

Depaftment  of  Tropical  Medicine 

Tulane  University 

New  Orleans  70112-2699 


ABSTRACT 

Pseudosuccinea  columella  (Sai/.  1817)  is  reported  for  the  first  time  from  the 
Dominican  Republic  and  from  the  island  of  Hispaniola.  It  occurs  together  with 
Fossaria  cuhensis  (Pfeiffer,  1839)  in  Puerto  Rico,  Cuba.  Jamaica,  and  Venezuela 
in  addition  to  the  Dominican  Republic  in  the  Caribbean  region.  The  two  lymnaeid 
species  can  be  differentiated  by  shell  and  animal  characteristics.  FascioiiasiH  is 
transmitted  in  this  region  p7-incipaUy  by  F.  cubensis,  but  the  occurrence  of  P.  col- 
umella in  some  countries  and  it.^  jtrobaJite  introduction  into  others  may  e.rlend  the 
range  of  this  disease. 


Pseudosuccinea  columella  (also  referred  to  as 
Lymnaea  columella  and  Lymnaea  (Pseudosuc- 
cinea) columella),  a  North,  Uentrtd  and  South 
American  species,  has  a[)parenUy  been  intro- 
duced into  certain  islands  of  the  Caribbean  and 
in  other  regions  such  as  Europe,  Africa, 
Australia  and  New  Zealand.  Maiek  and 
Chrosciechowski  (1964)  iind  Malek  and  Cogswell 
(1980)  reported  its  occurrence  in  Venezuela, 
Colombia,  Costa  Rica,  Pantima  and  Brazil  and 
reviewed  its  geogrtiphical  distribution. 
F'araense  (1982)  also  outlined  its  distribution  in 
the  neotropical  region. 


In  our  freshwater  snail  survey  in  the 
Dominican  Re{)ublic  we  encountered  P.  col- 
umella, but  it  was  less  common  (3  localities) 
than  the  other  lymnaeid,  Fossaria  cubensis 
(Pfeiffer,  1839)  (28  localities).  Previous  reports 
of  the  molluscan  fauna  of  the  Dominican 
Republic  did  not  include  P.  columella  (see 
Crosse,  1891),  nor  did  Alvarez  and  Cordero 
(1977)  in  their  work  on  fascioliasis  report  P.  col- 
umella; they  only  discussed  F.  cubensis. 

The  two  lymnaeid  species  can  be  differen- 
tiated morphologically  as  follows:  The  shell  of  P. 
columella  has  a  low  spire,  much  shorter  than  the 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      «7 


high  and  elongated  aperture,  with  a  large  and 
expanded  body  whorl.  The  surface  is  sculptured 
with  microscopic  raised  spiral  periostracal 
threads.  Lateral  teeth  of  the  radula  are  tri- 
cuspid. The  penis  sheath  is  short,  less  than  half 
the  length  of  the  prej)utium.  The  prostate  is  long 
and  narrowly,  roundly  cylindrical,  with  the 
proximal  end  somewhat  enlarged.  P.  rohonella 
is  aquatic,  with  a  tendency  to  be  amphibious. 

The  adult  shell  of  Fossarm  cubensis  is  small, 
with  well-impressed  sutures  and  usually  in- 
distinct spiral  lines.  The  spire  is  conic,  broadly 
pyramidal  or  acute,  the  aperture  is  much 
smaller  when  compared  to  that  of  P.  columella. 
The  columella  is  smooth;  the  inner  lip  of  the 
aperture  is  folded  backward,  forming  a 
moderate  or  broad  flat  expansion.  The  lateral 
teeth  of  the  radula  are  bicuspid.  The  penis 
sheath  is  two-thirds  the  length  of  the  preputium; 
the  prostate  is  ovate  and  rounded  at  the  ex- 
tremities. F.  eubensis  is  amphibious. 

Reports  of  P.  colitmelht  in  addition  to  F. 
cubensis  from  other  Caribbean  islands  are  from 
Puerto  Rico  (van  der  Schalie,  1948;  Harry  and 
Hubendick,  1964;  Leon-Dancel,  1970)  and  from 
Cuba  (Aguayo  and  Jaume,  1949).  Specimens 
received  from  Ms.  Charmine  Crawford  from 
Jamaica  are  also  P.  columella  and  F.  cubensis. 
The  two  species  are  also  known  to  occur  in 
Venezuela  (Malek  and  Chrosciechowski,  1964). 
P.  columella,  has  not  been  reported  from  St. 
Lucia  (Malek,  1965;  Barnish  el  ai.  1980), 
Guadeloupe  (Pointier,  1974,  1976),  Martiniciue 
(Guyard  and  Pointier,  1979),  or  Haiti  (Robart  et 
ai.  1976);  however,  F.  cubensis  is  present  in 
these  countries.  Pilsbry  (1910)  described  the 
subspecies  F.  citbensis  aspiraus  as  Liitunaea 
cubensis  aspirans  from  Barbados,  West  Indies. 

The  liver  tluke  Fasciola  hepatica.  which 
causes  fascioliasis  in  the  Americas,  occurs  in 
cattle  and  sheep-raising  countries,  parasitizing 
these  animals  and  other  herbivores.  Human 
fascioliasis  has  also  been  reported  from  alnwst 
every  American  country,  including  islands  of 
the  Caribbean.  There  are  several  early  reports 
of  high  prevalence  rates  on  Cuba  and  records  of 
some  human  cases.  There  are  also  records  of 
fascioliasis  in  the  Dominican  Republic  (Ueno  et 
ai,  1973;  Alvarez  and  Cordero,  1977),  Puerto 
Rico  (Frame  and  Bendezu,  1978;  Hillyer,  1981; 
Bendezu   et   a  I..    1982),    Martinique   (Gretillat, 


1966,  1967),  St.  Lucia  (Malek,  1965,  1980; 
Barnish  et  ai.  1980),  and  .Jamaica  (Anonymous, 
1960).  Malek  (1965)  signaled  the  occurrence  of 
fascioliasis  in  St.  Lucia  and  later  (1980)  gave 
details  of  prevalence  rates  in  several  estates  in 
the  north,  center  and  south  and  in  the  abattoir 
in  Castries,  the  capital.  Malek  (1965)  predicted 
the  occurrence  of  lymnaeid  snails  on  the  island 
and  shortly  after  the  report  was  written  F. 
cubensis  was  encountered  in  4  localities.  The 
snails  from  St.  Lucia  were  infected  exjierimen- 
tally  with  F.  hepatica  miracidia  of  St.  Lucian 
origin,  and  were  later  photographed  (Malek, 
1980).  F.  cubensis  from  St.  Lucia  was  also 
susceptible  to  the  schistosome,  Heterobilharzia 
aniericana  (Malek,  1967). 

From  all  reports  it  seems  that  in  the  Carib- 
bean region  the  principal  snail  intermediate  host 
of  F.  hepatica  is  F.  cubensis.  P.  columella, 
although  susceptible  to  experimental  infection 
(Leon-Dancel,  1970),  has  not  been  found 
naturally  infected  where  it  occurs  in  the  Carib- 
bean region,  although  natural  infection  has  been 
reported  in  Brazil  (Ueta,  1980).  There  is  also  a 
record  by  Van  Volkenberg  (1939)  (cited  in  Leon- 
Dancel,  1970)  and  one  by  Hillyer  (1981)  in  which 
P.  cdhanella  was  stated  to  be  an  intermediate 
host  in  Puerto  Rico.  The  natural  and  experimen- 
tal infections  in  P.  columella  indicate  that  its 
presence  in  some  Caribbean  countries,  and  its 
likely  introduction  in  others,  may  extend  the 
range  of  fascioliasis  in  the  region. 

References  in  the  literature  suggesting  that 
snails  other  than  lymnaeids,  for  example  Pliysa 
cuboixis  in  Cuba  or  terrestrial  snails,  can  act  as 
intermediate  hosts  for  F.  hepatica  are  er- 
roneous. Some  authors  tend  to  implicate  snails 
other  than  lymnaeids  as  transmitters  of 
fascioliasis  when  they  fail  to  locate  lymnaeids 
such  as  F.  cubensis  or  P.  cnhouclla  in  the  Carib- 
bean region.  Moreover,  Leon-Dancel  et  al. 
(1971)  tested  the  susceptibility  of  Physa  cubensis 
and  Aple.ra  marmorata  in  Puerto  Rico  and 
found  them  refractory  to  infection  with  F. 
hejiatica. 


LITERATURE  CITED 

Atrunv".  ('.  ('..  anil  .lauriie.  M.  L.  U)41).  Faniilia  Lyninaeidae. 
(in  i:i47-l'.l.'>:i)  '  'atiihyi,  ili  /.,s  Molliisms  ,!>■  Ciiha,  No.  4;t, 
Ciilia. 


68      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


Alvarez.  .1.  M.  V.  and  Conlcro.  E.  T.  C.  1977.  Fasciolosis 
bovina  en  la  Republica  Domiiiioana.  PuU.  Universidad 
Aulonoma S.  Domingo,  Vol.  224,  Col.  Cienciay  Techti.  No. 
5:1-189. 

Anonymous.  1960.  Annual  report  Ministry  Agriculture 
.lanuiica,  31  December  I9.')8. 

Barnish,  0..  Prentice,  M.  A.  and  Harris.  S.  1980.  Fasciola 
h<'i>(ittc<i  in  St.  Lucia,  West  Indies.  British  Vet.  Jour. 
136:299-300. 

Bendezu,  P.,  Frame.  A.  and  Hillyer,  G.  V.  1982.  Human 
fascioliasis  in  t'onrzal,  Puerto  Rico.  Jour.  Pnnisitol. 
68:297-299. 

Crosse,  H.  1891.  Faune  malacologique  terrestre  et  tluviatile 
de  I'ile  de  Saint  Domingue.  Jour.  Conrhyiiol.  39:69-211. 

Frame,  A.,  and  Bendezu,  P.  1978.  Bovine  fascioliasis  in 
Puerto  Rico.  Jour.  Parasitol.  64:136. 

Gretillat,  8.  1966.  Lymnaen  cuhensif!  Pfeiffcr  1911,  niol- 
lusque  bote  intermediaire  Ae  Fnsrioh  Iwpatirn  L.,  1758  a 
la  Martinique.  C.  R.  Acad.  Sri.  Pnris  262:2736-2739. 

1967.  Prospection  malacologique  aux  Antilles 

Francaises.  Observations  sur  I'ecologie  et  I'elevage  au 
laboratoire  de  Lymnapn  ruben.sis.  Ren  Elev.  Med.  Veter. 
Poys  Trop.  2:279-289. 

Ciuyard,  A.  and  Pointier,  .1.  I\  1979.  F^aune  malacologique 
<luIcaquicole  et  vecteurs  de  la  schistosomose  intestinale  en 
Martinii(ue  (Antilles  Francaises).  Arm.  Pitrnsitol.  (Paris) 
54:193-20.5. 

Harry,  H.  W.  and  Hubendick.  B.  1964.  Tlie  freshwater  pul- 
monale mollusca  of  Puerto  Rico,  Med.  Ootehorys  Mii.'i. 
Zooi  Ard.  136:1-77. 

Hillyer,  (!.  V.  1981.  Fascioliasis  in  Puerto  Rico.  [A  review.] 
Bol.  A  HOC.  Med.  Puerto  Rico  73:94-101. 

Leon-Dancel.  D.  1970.  Life  history  of  Lymnaeo  i-otioiielln 
(Say)  and  its  experimental  infection  with  Fri.sr/o/d  hepotico 
(L.).  Jour.  Agric.  Unir.  Puerto  Rieo  54:297-30.5. 

Ritchie,  L.,  and  Chiriboga.  .1.  1971.  Refractive- 

ness  of  P/ij/.S((  ruherma  (Pfeiffer)  and  .Aple.rn  mormorntn 
(Guilding)  to  Fa.friota  hepotico  (L.).  Jour.  .Agric.  Unir. 
Puerto  Rico  55:267-270. 


Malek.  F>.  A.  1965.  Freshwater  and  land  snails  of  St.  Lucia, 

the  West  Indies.  Arner.  Malncol.  Union.  Annual  Reports, 

p.  38. 
1967.  E^xperiniental  infection  of  several  lym- 

naeid    snails    with    Hetero))ithnrzi(i    nmi'ricono.    Jour. 

Pariiiiitol.  53:700-702. 
1980.  Snail   Trnnaniitted  Paraaitic  Diseases, 

Vol.  n,  CRC  Press,  Boca  Raton,  Florida.  324  pp. 
and  Chroscieehowski,  P.  1964.  Lymnaea  (Pseu- 


dosuccinea)  columella  from  Venezuela,  and  notes  on  distri- 
bution <){  P.'ieudosuecinea.  The  Nautilus  78:54-56. 

and  Cogswell,  F.  B.  1980.  Lymnaea  (Pseudosuc.- 


cinea)  columella  in  Colombia.  The  Nautilus  94:112-114. 
Paraense,  W.  L.  1982.  Lymnaea  riatrix  and  Lymnaea  col- 
umella in  the  Neotropical  region:  A  distributional  outline. 

Mem.  Inst.  Clswaldo  Cruz  77:181-188. 
Pilsbry,  H.  A.  1910.  Lymnaea  cubensis  nspirans  n.  subsp. 

The  Nautilus  23:\2{)'. 
Pointier,  .1.  P.  1974.  F^iune  malacologique  dulcaquicole  de 

I'ile  de  la  (Hiadeloupe  (Antilles  Francaises).  Bull.  Mus. 

Hist.  Natur..  Paris,  3rd  Ser.,  No.  235,  Zool.,  1.59:905-933. 
1976.  Repartition  locale  el  biogeographie  des 

mollusques    dulcaquicoles    de    la    Guadeloupe    (Antilles 

Francai.ses).  Malacol.  /?e<'i>H' 9:85-103. 
Robart,  G..  Mandahl-Barth.  G.  and  Ripert.  C.  1976.  Inven- 

taire,  repartition  geographique  et  ecologie  iles  mollusques 

dulcaquicoles  d'Haiti  (Caraibes).  Haliotis  8:159-171. 
Heno,  H..  Alvarez.  .J.  M.  V.,  Mergen.  A.  M.  R.  de  and 

Sanchez,  V.  M.  1973.  Observation  on  the  prevalence  of 

parasitic  diseases  in  cattle  especially  fascioliasis,  in  the 

Dominican  Republic.  Nat.  Inst.  Animal  Hlth.  Quarterly 

(Tokyo)  13:59-68. 
Ueta,  M.  T.  1980.  Ocorrencia  de  infecvao  natural  dv  Faxciola 

hepatica  Linnaeus.  1758  em  Lymnaea  columella  Say.  1817 

no  Vale  do  Paraiba.  SP.  Brasil.  Rec.  Saude  Publico  14: 

230-233. 
van  der  Schalie.  H.  194cS.  The  land  and  fresh  water  niollusks 

of  Puerto  Rico.  Misc.  Publ.  Mus.  Zool.  Univ.  Michigan, 

No.  70. 


ANNOUNCING  THE  CO. A. 
19S6  CONVENTION 

The  Concholo^ists  of  America's  14th  annual 
convention  will  he  held  at  the  Sheraton  \'ankt'e 
Trader  Hotel  at  Fort  I  Lauderdale,  Florida,  from 
July  1')  throujjjh  July  19,  198(5.  The  convention 
will  he  hosted  hy  the  Broward  Shell  Cluh,  and 
the  CO. A.  welcomes  all  shell  collectors, 
whether  novice  or  advanced,  as  well  as  shell 
dealers  and  scientists  from  aroinid  the  country 
and  the  world. 

For  further  information  and  pre-registration 
packets,  please  contact  Ruth  ('hesler.  Conven- 
tion Chairman,  at  7401  S.  W.  7th  Street.  Planta- 
tion, Florida  8:5317. 


NEWS 

JULY  1986  AMU  MEETING 
The  American  Malacological  Union  and  the 
Western  Society  of  Malacolojjists  will  have  a 
joint  meetin,ti;  from  July  1-5  at  the  Sheraton 
Hotel  in  the  heart  of  historical  Monterey,  Calif. 
The  three  symposia  will  be  on  the  biolojjy  of 
opisthohranchs,  life  history  of  cephalopods  and 
molluscan  morphological  analyses.  Housing, 
travel  and  registration  details,  already  sent  to 
AMU  members,  may  also  be  obtained  from 
President  Dr.  James  Nybakken,  Marine  Labora- 
tories. P.O.  Box  450,  Moss  Landing,  CA 
95039-0450. 


Vol.  100(2)  April  ::5(),  1986  THE  NAUTILUS      69 

SOME  ASPECTS  OF  THE  EVOLUTION  OF  THE  RUDIST  PELECYPODS 


David  Nicol 

Box  14376,  University  Station 
Gainesville,  FL  32604 

ABSTRACT 

The  evolution  of  the  rudist  pelecypods  was  remarkably  rapid  even  as  compared 
to  many  mammalian  groups.  One  new  genus  appeared  every  1.900,000  years  in 
the  Late  Jurassic,  hi  the  Early  Cretaceous,  one  neiv  genus  appeared  every 
1,370,000  years,  hut  iii  the  Late  Cretaceous  the  rate  of  evolution  accelerated  great- 
ly when  one  new  genus  appeared  every  3^5,000  years.  For  the  entire  time-span  for 
the  existence  of  the  rudists,  one  new  genus  appeared  every  738,000  years.  Like  the 
dinosaurs,  the  first  rudists  were  of  modest  size.  Beginning  in  the  AUiian  stage, 
some  rudists  attained  a  size  as  large  as  any  pelecypods  that  have  ever  lived. 


Some  additional  aspects  of  the  evolution  of  the 
rudists  are  given  here  because  they  are  either 
not  mentioned  or  inadequately  covered  in  a 
manuscript  now  in  press  (D.  S.  Jones  and  D. 
Nicol:  Origination,  survivorship,  and  extinction 
of  rudist  taxa).  Some  of  the  data  used  herein 
were  updated  by  examining  the  Zoological 
Record  from  1966  to  1981.  The  main  source  of 
data  is  the  Treatise  on  Invertebrate  Paleon- 
tology (C.  Dechaseaux  and  A.  H.  Coogan,  1969). 

The  aspect  that  is  particularly  emphasized 
here  is  the  rapid  increase  in  diversity  at  the 
generic  level,  especially  during  the  Late 
Cretaceous  or  the  last  29  million  years  of  the  ex- 
istence of  the  rudists.  The  15  Jurassic  and 
Cretaceous  stages  listed  in  Table  1  are  based 
primarily  on  ammonite  zones  and  are  not  all  of 
the  same  duration.  The  duration  in  years  of  each 
stage  was  taken  from  Palmer,  1983.  The  15 
stages  can  be  subdivided  into  three  groups  for 
studying  the  changes  of  rates  of  evolution:  the 
three  Jurassic  stages  (Oxfordian  through  Titho- 
nian),  the  Early  Cretaceous  stages  (Berriasian 
through  Albian),  and  the  Late  Cretaceous 
stages  (Cenomanian  through  Maastrichtian). 

Rudist  evolution  began  at  a  rather  modest 
rate  in  the  Late  Jurassic  with  a  new  genus  ap- 
pearing approximately  every  1,900,000  years. 
The  evolutionary  rate  did  not  accelerate  greatly 
during  Early  Cretaceous  time  when  a  new 
genus  appeared  every  1,370,000  years.  How- 
ever, beginning  with  the  Cenomanian,  the 
earliest  stage  of  the  Late  Cretaceous,  evolution 
speeded  up  greatly,  and  a  new  genus  arrived 
every  345,000  years.  The  evolutionary  rate  for 


TABLE  1.  The  I.t  Jurassic  and  Cretaceous  stages  with 
their  duration  in  millions  of  years,  number  of  new  genera  per 
stage,  and  new  genera  per  year.  Oxfordian -Tithonian.  Late 
Jurassic;  Berriasian -Albian,  Early  Cretaceous;  Ceno- 
manian- Maastrichtian,  Late  Cretaceous. 


stages 

Duration 

Mew 

:e^  £-n-rd 

in  years 

jenera 

per  ye^r 

■■■'aastrichtia.a 

",000,000 

17 

470,000 

Campanian 

9,500,000 

13 

730,000 

Santonian 

2,500,000 

15 

100,000 

Coniacian 

1,000,000 

10 

100,000 

Turonian 

2,500,000 

13 

192,000 

Cenomanian 

5,500,000 

lo 

344,000 

Albian 

15,500,000 

10 

1 

, 55o,000 

Aptian 

0,000,000 

7 

"57,000 

Barresnian 

5,000,000 

9 

025,000 

Hauterivian 

7,000,000 

0 

— 0— 

Valanginian 

7,000,000 

5 

1 

,400,000 

rerriasian 

0,000,000 

4 

1 

,500,000 

Tithonian 

8,000,000 

4 

2 

,000,000 

Kimmeridgian 

4,000,000 

0 

— 0— 

Oxfordian 

7,000,000 

0 

1 

,107,000 

Totals 

94,500,000 

12'! 

73 ",000 

the  total  15  stages,  or  the  complete  life  span  of 
the  rudists,  is  the  appearance  of  a  new  genus 
every  738,000  years.  This  would  be  considered 
exceedingly  rapid  evolution  amongst  the  pelecy- 
pods in  a  stable  marine  environment  and  is  com- 
parable  to   rapid  evolutionary   rates   in   many 


THE  NAUTILUS 


April  ;i(),  1986 


Vol.  100(2) 


groups  of  mammals. 

The  rapid  acceleration  of  evolution  of  the 
rudists  during  the  Late  Cretaceous  appears  to 
be  based  on  two  phenomena.  First,  the  three 
larger  and  more  rapidly  evolving  families 
(Caprinidae,  Radiolitidae,  and  Hippuritidae) 
either  did  not  appear  until  the  Late  Cretaceous 
or  did  not  become  diverse  until  that  time.  Sec- 
ond, the  rudist  faunas  became  more  provincial 
and  ecologically  complex  in  the  Late  Cretace- 
ous. Endemism  of  the  rudist  genera  increased 
greatly  in  the  Late  Cretaceous,  and  the  rudist 
communities  were  generically  most  diverse  in 
the  Caribbean  region  during  the  Maastrichtian 
(Kauffman  and  Sohl,  1974). 

As  more  paleontological  study  is  done  in  the 
more  remote  areas  with  Jurassic  and  Cretace- 
ous rocks  in  the  Tethyan  realm,  a  greater 
number  of  new  genera  discovered  will  be  in  the 
Late  Cretaceous  rather  than  in  the  Early 
Cretaceous  and  Late  Jurassic. 

In  the  rates  of  evolution  within  the  L5  stages 
in  Table  1,  it  is  worthwhile  to  point  out  a  few 
additional  features.  After  a  good  start  in  the  Ox- 
fordian  Jurassic,  the  evolution  of  the  rudists 
slowed  somewhat  with  a  gap  of  no  new  genera 
in  the  Kimmeridgian,  a  time  span  of  four  million 
years.  This  slowing  of  the  rate  of  evolution  con- 
tinued into  the  first  three  stages  of  the  Early 
Cretaceous  with  a  climax  in  the  Hauterivian 
when,  again,  no  new  genera  have  so  far  been 
discovered.  The  unusual  aspect  of  the  Hauteri- 
vian gap  is  that  it  encompassed  approximately 
seven  million  years. 

Beginning  in  the  Barremian,  evolution  accel- 
erated greatly  and  this  may  have  been  caused 
partially  by  the  onset  of  the  two  most  diverse 
families,  the  Caprinidae  and  Radiolitidae.  The 
rate  of  evolution  then  slowed  in  the  next  two 
stages  which  end  in  Early  Cretaceous  times. 
New  genera  appeared  with  exceeding  rapidity 
in  the  Cenomanian  and  continued  unabated  until 
the  Campanian  and  then  accelerated  again  in 
the  Maastrichtian.  In  the  latest  Cretaceous,  new 
rudist  genera  were  still  appearing  at  a  very 
rapid  rate. 

Although  extinction  of  genera  occurred  rapid- 
ly in  the  Late  Cretaceous,  the  large  number  of 
new  genera  in  each  stage  tended,  with  some  ex- 
ceptions, to  increase  the  total  number  of  genera 
in   each   stage   to   a   maximum   of  43   in   the 


Maastrichtian.  This  total  of  43  includes  the 
holdovers  from  earlier  stages  as  well  as  the  new 
arrivals.  The  present  total  of  rudist  genera  is 
128  so  that  within  a  period  of  no  more  than  eight 
million  years,  one  third  of  all  the  rudist  genera 
became  extinct.  Five  of  the  seven  rudist  families 
also  became  extinct  in  the  Maastrichtian. 

The  decline  and  extinction  of  the  rudists  oc- 
curred in  slightly  less  than  8.5%  of  their  total 
geologic  history.  This  sudden  demise  of  the 
rudists  at  their  zenith  at  the  end  of  the 
Cretaceous  is  unlike  the  extinctions  of  the 
trilobites,  graptolites,  conodonts,  and  rostro- 
conchs  where  there  was  a  long  period  of  decline 
in  diversity  before  extinction.  Even  the  am- 
monites and  dinosaurs  experienced  a  longer 
period  of  decline  than  the  rudists.  It  is  most  dif- 
ficult and  highly  speculative  to  explain  such  a 
sudden  extinction  in  a  group  of  marine  pelecy- 
pods  that  had  been  so  highly  successful. 

The  rudist  pelecypods,  like  the  dinosaurs, 
began  with  small-sized  species.  Some  of  the 
early  rudists  were  about  20  mm  long,  and  small- 
sized  rudists  are  also  found  at  the  end  of  the 
Cretaceous  in  Maastrichtian  strata.  Particularly 
among  the  diverse  families  Caprinidae  and 
Radiolitidae,  some  stocks  attained  large  size  as 
early  as  the  Albian,  and  large  species  continued 
to  occur  into  the  Maastrichtian.  However,  even 
by  Albian  time,  50  million  years  had  passed, 
which  is  more  than  half  of  the  total  time  span  of 
the  rudists.  Titanosarcolites  attained  a  length  of 
more  than  2  m,  Bournonia  reached  a  diameter 
of  more  than  0.6  m,  and  the  shell  wall  of 
Durania  attained  a  thickness  of  at  least  10  cm 
(B.  F.  Perkins,  1969,  p.  N751).  These  large 
rudists  were  among  the  largest  pelecypods  that 
have  ever  lived. 

Acknowledgments 

I  am  indebted  to  Gary  S.  Morgan  of  the 
Florida  State  Museum  for  making  some  helpful 
suggestions  that  improved  this  paper. 

LITERATURE  CITED 

Ilechaseaux.  ('.,  and  A.  H.  Coogan.  1969.  Hippuritacca- 
Systeniatic  descriptions,  p.  N776-817.  In  R.  C.  Moore  (ed.). 
Treatise  on  Invertebrate  Paleontology,  Part  N,  Vol.  2, 
Mollusca  6,  Bivalvia.  Univ.  Kansas  Press,  Lawrence. 

Jones,  D.  S.,  and  D.  Nicol.  Origination,  survivorship,  and 
extinction  of  rudist  taxa.  Jour.  Paleo.  60:107-115. 

Kauffman.  E.  ti..  and  N.  F.  Sohl.  1974.  Structure  and  evolu- 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      71 


tion  of  Antillean  Cretaceous  rudist  frameworks.  Verhand- 
lungen  Naturfor.'ichende  Gesellschaft.  Basel.  84:399-467. 

Palmer,  A.  R.  198,3.  The  decade  of  North  American  geologT,- 
1983  geologic  time  scale.  Geology  ll:.503-504. 

Perkins.   B.   F.   1969.   Rudist   morphology' -Shell  size  ami 


shape,  p.  N7.")l.  In  R.  C.  Moore  (ed.).  Trcntixc  on  hirci-tr- 
bmte  Paleontology,  Part  N,  Vol.  2,  Mollusca  6,  Bivalvia. 
Univ.  Kansas  Press,  Lawrence. 
Zoological  Record.    1966-1981.   The  Zoological   Society  of 
London. 


A  NEW  SOMATOGYRUS  FROM  THE  SOUTHWESTERN  OZARKS  WITH 

A  BRIEF  REVIEW  OF  THE  HYDROBHDAE  FROM  THE 

INTERIOR  HIGHLANDS  (GASTROPODA:  PROSOBRANCHIA) 


Mark  E.  Gordon 

Department  of  Zoology 
University  of  Arkansas 
Fayette vi"lle,  AR  72701 

ABSTRACT 

A  previously  unknown  Somatogyrus  is  described  from  the  Elk  River,  McDoridld 
County,  Missouri.  With  this  description,  the  number  of  Hydrobiidae  presently 
recorded  from  the  Interior  Highlands  increases  to  15,  including  11  regional 
endemics.  Nine  species  are  endemic  to  the  Ozark  Plateaus,  while  two 
Somatogyrus  are  restricted  to  the  Ouachita  Mountains.  Fire  of  the  Ozark 
endemics  are  obligate  troglohites.  Troglobitic  Amnicola  antroecetes  (Hubrichf)  is 
considered  distinct  from  epigean  A.  aldrichi  (Call  and  Beecher). 


Excepting  the  Unionoidea,  little  study  has 
been  afforded  the  aquatic  Mollusca  of  the  In- 
terior Highlands.  Among  the  poorer  known 
groups,  regional  accounts  of  the  Hydrobiidae 
are  rare.  The  primary  sources  of  pulilished 
records  have  been  original  descriptions  of 
several  endemic  species  (e.g.  Walker,  1915; 
Hinkley,  1915;  Hubricht,  1979),  although  the  oc- 
casional collection  of  hydrobiids  has  been  noted 
in  a  few  species  surveys  (e.g.  Sampson,  1913; 
Wheeler,  1918;  Gordon,  1981).  A  recent  faunal 
survey  of  drainage  basins  in  southwestern 
Missouri  and  northwestern  Arkansas  (Gordon, 
1980)  recovered  several  species  of  Hydrobiidae. 
A  previously  unknown  Somatogyrus.  collected 
from  the  Elk  River,  McDonald  County, 
Missouri,  is  here  described. 

Family  Hydrobiidae  Troschel  1857 
Subfamily  Lithoglyphinae  Fischer  1885 
Somatogyrus  rosewateri  new  species 
Shell    descripfiori     of   holotype-SheW    sub- 


globose,  medium  size  for  Somatogyrus.  solid, 
dextral,  4.25  whorls,  turbinate,  subhyaline;  body 
whorl  .90  times  axial  height,  greatly  inflated; 
[penultimate  whorl  .34  times  width  of  body  whorl 
(Fig.  1;  see  Table  1  for  holotype  measurements 
and  range  of  paratypes).  Spire  very  short, 
depressed,  broadly  conic;  sutures  moderately 
impressed;  protoconch  flattened  (Fig.  2). 
Periostracum  greenish  (may  be  masked  by  color 
of  visceral  mass  showing  through  shell)  with 
shallow,  oblique  growth  striae;  protoconch  or- 
namented with  fine,  spiral  ridges  (Fig.  3).  Aper- 
ture broadly  ovate,  .74  times  axial  height,  .80 
times  height  and  .55  times  width  of  body  whorl, 
plane  of  aperture  at  30°  to  shell  axis.  Peristome 
complete  across  parietal  wall  by  a  callus  which 
continues  and  thickens  across  columella  (callus 
in  some  individuals  may  be  slightly  reflected 
over  columella,  partially  obscuring  umbilicus); 
lip  straight  along  parietal  wall,  curving  con- 
cavely  near  the  umbilicus  and  along  columella, 
parietal-columellar  junction  non-angular.  In 
lateral  view  (Fig.  4),  periphery  of  parietal  lip 


72      THE  NAUTILUS 


April  30,  1986 


Vol.  100  (2) 


FIGS.  1-5.  Suiniit(i(j!irus  nuiruvlcn  n.  .sp.  1,  apei'tural  view 
(if  paratype  (3.9  x  3.8  mm.);  2,  juvenile  paratype;  3,  suture 
and  pr<)t(jconeh  sculpture  iif  paratype.  The  liarely  visible 
white  bar  in  the  upper  left  is  10  micrcms  in  length.  4,  lateral 
view  of  paratype  (4.2  x  3.8  nun.);  5,  urnhilical  view  (if 
paratype  (shell,  3.fi  x  3.5  mm.). 


approximately  parallel  to  plane  of  aperture,  col- 
umella straight  but  recedes  towards  base,  and 
parietal-columellar  junction  forma  an  obtuse 
angle.  Umbilicus  relativi'ly  open  for  Somnfo- 
gyrus  (Fig.  5);  passes  into  a  shallow,  canal-like 
depression  continguous  to  the  columellar  Iif). 
Basal  lip  slightly  receded  due  to  angle  of  aper- 
ture; area  of  base-columcllar  junction  projects 
forward  about  ecjual  to  the  distance  of  the  basal 
lip  recession  (Fig.  4). 

Operculum  chitinous,  ovate,  horn-colored; 
paucispiral,  consisting  of  about  three  whorls; 
nucleus    subcentral,    located    slightly    left    of 


midline  in  lower  third;  outer  surface  sculptured 
with  many  fine  growth  striae,  peripheral  margin 
without  striae  and  edge  appears  slightly  ragged 
(Fig.  6). 

Soft  paW.s- Living  animals  translucent  (odon- 
tophore  complex  visible  through  proboscis), 
whitish.  Mantle  collar  and  dorsal  aspects  of  pro- 
boscis, head,  tentacles,  and  foot  speckled  with 
sub-epidermal  gold  to  orangish  gold  chromato- 
cytes  (Fig.  7);  dorsal  surface  and  V3  thickness  of 
visceral  mass  infused  with  golden  color.  Dense 
concentrations  of  melanin  at  base  of  tentacles, 
extends  dorso-medially  in  front  of  eyes  a  short 
distance  and  posteriorly  on  to  dorsum  of  head 
(Fig.  8);  a  narrow  line  of  melanin  runs  laterally 
from  in  front  of  eye  forward  about  V4  to  Vs 
length  of  tentacle;  dorsal  and  lateral  surfaces  of 
mantle  with  mottled  patterns  of  melanin  (Fig. 
7),  35%  of  paratypes  (n  =  52)  lacked  mantle 
melanin.  Shells  of  living  snails  appear  amber  to 
orangish  due  to  visceral  and  mantle  pigmenta- 
tion showing  through  shell,  darkens  to  brown  in 
spire;  melanistic  snails  have  slightly  browner 
tint.  Penis  unpigmented.  Preserved  animal 
white. 

Foot  broad  (Fig.  7);  anterior  truncated,  ex- 
tending forward  to  eyes;  posterior  tapers  to  a 
rounded  point.  Head  (Fig.  8)  extends  anterior  of 
shell  to  expose  eyes;  proboscis  relatively  long 
and  slender,  curved  ventrally;  tentacles  long 
and  slender,  eyes  positioned  ventro-laterally  at 
base.  Penis  (Fig.  9)  slender,  simple;  tapers  to  a 
fine,  blunt  point;  dorso-ventrally  flattened,  dor- 
sum slightly  convex,  venter  flatly  concave;  vas 
deferens  positioned  near  posterior  margin. 
Penis  arises  a  short  distance  inside  mantle  cavi- 
ty above  and  behind  the  right  tentacle,  tends  to 
curve  counter  clockwise;  when  extruded, 
emerges  from  shell  directly  behind  right  ten- 
tacle and  extends  below  it  (Fig.  8). 

Type  locality -M\ssour\,  McDonald  County, 
Elk  River  at  Missouri  Department  of  Conserva- 
tion Mt.  Shira  Public  Fishing  Access,  1.6  km 
south  of  Ginger  Blue  (R33W,  T21N,  sections  1 
and  2).  Holotype:  University  of  Colorado 
Museum  of  Zool()g>'  (UCM)  no.  32812;  collected 
24  October  1985  by  Mark  E.  Gordon.  Allotype: 
UCM  no.  32813;  same  data  as  holotype.  Para- 
types: ten  specimens  each  deposited  at  UCM, 
U.S.  National  Museum  of  Natural  History, 
Academy  of  Natural  Sciences  of  Philadelphia, 


Vol.  100(2) 


A|:)ril  80,  1986 


THE  NAUTILUS      73 


TABLE  1.  Dimensions  of  holotype,  allotype,  and  paratypes  oi'  SumiifiHii/nis  niKPimferi  (measurements  in  mm):  H  = 
height,  W=  width,  BH  =  body  whorl  height,  PW  =  penultimate  whorl  width,  A  =  aperture,  AH  =  aperture  height, 
AW  =  aperture  width. 


H 

W 

W/H 

BH     BH/H 

W/BH 

PW 

PW/W 

AH 

AW     AW/AH 

Plane  of  A 

Holotype 

(9i 

J. 9 

i.8 

.97 

3,5 

90 

1.09 

1.  3 

.34 

2.8 

2.1 

75 

30° 

Allotype 

(o-) 

3.4 

3.4 

1.00 

3.1 

91 

1.10 

1.0 

.29 

2.5 

1.9 

76 



Paratype 

3.7 

3.6 

.97 

3.2 

86 

1.13 

1.2 

.33 

2.3 

2.1 

91 



„„ 

4.0 

3.7 

.93 

3.5 

88 

1.06 

1.2 

.32 

2.7 

1.7 

63 

34° 

„„ 

4.1 

3.9 

.95 

3.6 

88 

1.08 

1.2 

.31 

2.8 

2.4 

86 

29° 

„„ 

4.0 

3.8 

.95 

3.6 

90 

1.06 

1.1 

.29 

2.9 

2.2 

/6 



tin 

3.9 

3.7 

.95 

3.4 

87 

1.09 

i.2 

.32 

2.5 

2.1 

84 

34° 

II  It 

3.8 

3.8 

l.OU 

3.1 

89 

1.12 

1.3 

.34 

2.4 

2.1 

88 

35° 

„„ 

3.8 

3.5 

.92 

3.4 

89 

1.03 

1.3 

.3/ 

2.5 

2.1 

84 

30° 

(Fig. 

1) 

3.9 

3.8 

.97 

3.5 

90 

1.09 

1.1 

.29 

2.5 

2.1 

84 

30° 

(Fig. 

3,4) 

4.2 

3.8 

.90 

J. 7 

88 

1.03 

1.3 

.34 

2.8 

2.2 

79 

30° 

(Fig. 

5) 

3.6 

3.5 

.97 

3.3 

92 

1.06 

1.0 

.29 

2.5 

1.9 

76 

27° 

FIGS.  6-9.  Somatogyrus  rosewateri  n.  sp.  6,  operculum;  7,  animal  extruded  from  shell 
(composite  drawing  from  live  and  rela,\ed  specimens;  body  slightly  distended  from  relaxa- 
tion with  sodium  pentobarbital):  C  =  chromatocytes,  E  =  eye,  F  =  foot,  M  =  melanin. 
Mm  =  mantle  melanin  showing  through  shell.  Mo  =  mouth,  0  =  operculum,  P  =  proboscis, 
T  =  tentacle;  8,  dorsal  view  of  head:  Pn  =  penis;  9,  penis:  V  =  vas  deferens. 


Museum  of  Comparative  Zoology,  University  of 
Michigan    Museum    of    Zoology,     and    FieW 
Museum;  same  data  as  holotype. 
Distribution -Found  only  in  the  middle  sec- 


tion of  Elk  River  (Fig.  10).  To  date,  this  species 
has  been  collected  from  only  the  type  locality 
and  below  an  old  mill  dam  at  Noel,  McDonald 
County,  Missouri.  It  was  not  recovered  from 


74      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


ARIANSAS 

FKi.  Kl.  Distriliution  of  Siini.atofiyrus  riiaeiratcri  (sdlid 
triangle  is  the  type  locality). 

areas  upstream  of  the  type  locality  nor  the  river 
channel  downstream  from  the  Missouri- 
Oklahoma  border  which  has  been  inundated  by  a 
reservoir  on  the  Neosho  River  (Grand  Lake  0' 
the  Cherokees).  The  known  range  of  this  species 
occurs  in  the  Springfield  Plateau  region  of  the 
Ozarks. 

//rr6;7a/- Specimens  were  collected  at  the 
type  locality  from  a  series  of  riffles.  Substrate 
was  composed  of  gravel  and  cobble-sized 
materials  (mainly  chert).  At  Noel,  habitat  was 
below  the  outfall  of  a  small  mill  dam.  Substrates 
were  primarily  stepped  and  broken  bedrock 
with  some  cobble.  Gradients  at  both  sites  were 
fairly  high  and  water  quality  was  quite  good 
(Table  2). 

Efyyuology -Th\i^  s|)ecies  is  named  in  memory 
of  Dr.  Joseph  Rosewater,  late  Curator  of 
Mollusks,  U.S.  National  Museum  of  Natural 
History. 

Discussion 

The  comparative  difference  between  S. 
rosewateri  and  previously  described  species  in 
the  genus  are  shown  in  Table  3. 


TABLE  2.  Physicd-cheniical  nieasurenierits  madi 
locality.  Elk  River;  20  November,  1979. 


at  type 


Water  temperature 

Tiirliiility 

Specific  coruiuctance 

Alkalinity 

I'll 

Nitrate 

Color 


15°C 

1.6  NTU 

2(j0  fjmhos/cm 

140  mfT/l  total  CaCUj 

8.1.-) 

1..')  mg/l 

neglij^ble 


Besides  Somatoyyru,^  rosewateri.  the  only 
other  hydrobioids  collected  from  the  Elk  River 
basin  were  the  nymphophiline  Cincinnatia  in- 
tegra  (Say)  and  the  pomatiopsid  Pomatiopsi>i 
lapidaria  (Say)  (Gordon,  1980).  Two  other 
Somatogyrus  are  known  from  the  Ozark 
Plateaux.  Somatogyrus  depressus  (Tryon),  an  in- 
habitant of  the  upper  Mississippi  River  valley, 
occurs  in  the  Osage  River,  central  Missouri 
(Thompson,  1984)  and  S.  crassilahris  Walker, 
an  Ozark  endemic,  is  apparently  known  only 
from  its  original  collection  from  the  North  Fork 
of  the  White  River,  Arkansas.  In  discussing 
phylogenetic  relationships  among  North  Ameri- 
can Lithoglyphinae,  Thompson  (1984)  noted  dif- 
ficulties in  pursuing  such  studies  due  to  a 
relatively  large  number  of  undescribed  species 
and  a  paucity  of  morphological  data  for  de- 
scribed species.  For  Somatogyrus.  he  did  list 
several  characters  useful  for  specific  com- 
parisons. Available  data  for  the  three  Ozark 
species  and  two  Ouachita  Mountains  endemics 
(Table  3)  suggests  that  the  Ozark  species  may  be 
fairly  closely  related.  Walker  (1915)  also  con- 
sidered S.  wheeler i  Walker  closer  to  S. 
crassilahris  than  to  its  Ouachitan  congener. 
However,  both  Ouachita  Mountains  endemics 
may  have  evolved  from  other  lineages  (e.g. 
Alabama  River).  Considerably  more  information 
is  necessary  before  such  speculative, 
phylogenetic  relationships  may  be  confirmed.  In 
some  instances,  complete  analyses  may  no 
longer  be  possible.  Many  hydrobiids,  such  as  S. 
era.ssilabrls.  S.  amnicoloides  Walker,  and  S. 
wheeleri.  are  known  only  from  their  type 
localities.  Habitat  alterations,  such  as  reservoir 
construction,  may  have  been  deleterious  for 
species  with  such  highly  restricted  distributions. 

Fifteen  species  of  Hydrobiidae  have  been 
documented  from  the  Interior  Highlands  (Table 
4).  Four  are  wide-ranging  species.  In  the  case  of 
Prohythvnella  lacu.'^tris  (Baker)  (mistakenly 
listed  as  F.  binneyana  (Hannibal)  in  Gordon, 
1981)  and  Somutogyrus  depres.'<y.s.  their  south- 
ern distributional  limits  occur  in  the  Ozarks. 
Clncinnaila  integra  and  Birgella  subglohosa 
(Say)  have  considerably  larger  ranges  than  the 
previous,  more  northerly  distributed  species 
(see  Rurch  and  Tottenham,  1980;  Thompson, 
1984).  The  remaining  eleven  species  are 
endemic  to  the  Interior  Highlands.  Somatogyrus 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      75 


TABLE  3.  Variation  in  shell  morpholo^  among  Interior  Highlands  Somalomirti^  (data  adapteil 
from  Baker,  1928;  Burch  and  Tottenham,  1980:  Hinkley.  1915;  Walker,  191.');  and  tliis  study). 

depress  us   crassilabris    rosewateri  wheeleri   ainnicoloides 


Shell    obesity    (W/H)  .78-1.00 

Size 

medium    {<4   imi)(0)  0 

small    (>  4   mm)  ( 1) 

Spire 

depressed  (>15%  H)(0)    0 
elevated  (<15%  H)  (1) 

Protoconch 

flattened  (0)  0 

elevated  (1) 

Par ietal-colume liar 
wall  junction  of  lip 
smooth  curve  ( 0 )         0 
angular  (1) 

Umbilicus 

narrow  { 0 )  0 

open  ( 1 ) 

Widest  expansion 
of  outer  lip  periphery 
above  mid-height  (0)     0 
below  mid-height  (1) 

Aperture  apex  relation 
to  parietal  wall 
fused  (0)  0 

detached  (1) 


.77-. 92 


.90-1.00 


.93 


amnicoloides  and  S.  ivheeleri  are  presently  the 
only  endemic  hydrobiids  that  have  been  iden- 
tified from  the  Ouachita  Mountains  province. 
Four  epigean  and  five  troglobitic  species  are 
endemic  to  the  Ozark  Plateaus.  The  maj(jrity 
(eight)  are  restricted  to  the  eastern  half  of  the 
province.  The  only  endemic,  western  Ozark 
hydrobiid  presently  recognized  is  S.  ro^tewaferi: 
however,  a  single  specimen  of  Somntogyrus  has 
been  collected  from  the  headwaters  area  of  the 
White  River,  Arkansas  (Gordon,  1980).  This 
specimen  was  too  eroded  to  allow  identification 
beyond  genus  and  sulisequent  collecting  failed 
to  recover  additional  specimens. 

Most  of  the  Ozark  endemic,  obligate  troglobi- 
tic Hydrobiidae  have  been  referred  to  Amnicoli- 
nae:  Anniicoln.  Burch  and  Tottenham  (1980) 
noted  that  this  classification  was  tentative  due 
to  the  lack  of  anatomical  flata  for  these  species. 


However,  Antrobia  culreri  Hubricht  was  rele- 
gated to  the  Lithoglyphinae  without  justifi- 
cation by  Burch  and  Tottenham  (1980).  The 
anatomy  of  this  species  remains  all  but 
unknown.  The  brief  description  of  the  radula 
presented  by  Hubricht  (1971)  does  not  appear  to 
meet  the  radular  criterion  for  Lithoglyphinae 
established  by  Thom()son  (1984)  and  differs 
from  that  of  Amniroln  only  by  the  size  of  the 
central  tooth  mesocone.  Hubricht  (1971) 
described  the  penis  of  Antrohid  as  "simple, 
tapering  to  a  point,  without  appendages."  This 
morphology  is  characteristic  of  the  lithoglyphine 
penis  (Thompson,  1984)  and  is  prolialily  the 
basis  for  the  classification  in  Burch  and  Totten- 
ham (1980).  Unfortunately,  Hubricht  (1971)  did 
not  describe  the  number  of  ducts  penetrating 
the  (lenis.  Thompson  (1984)  considered  the 
Amnicolinae,  with  two  penial  ducts,  remote  in 


76      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


TABLE  4.  Hydrobiidae  known  from  the  Interior  Highlands. 

Hydrobiinae 

Prohythinctta  iacu.stritt  {Baker,  1928) 
Lithoglyphinae 

Som<itogy7-us  amnicoloides  Walker,  1915 

Su7n.ato(fi/rus  crassilabris  Walker,  1915 

Somntogi/rus  depressus  (Tryon,  1862) 

Somntotfynis  rosewateri  new  species 

Somatog-yruit  wheeleri  Walker,  1915 
Nymphophilinae 

Birgeila  subglohona  (Say,  1825) 

Cincinnalia  Integra  (Say,  1821) 

Marstonia  ozarkensis  (Hinkley,  1915)' 
Amnicolinae 

Amnicolii  aid  rich  i  (Call  and  Beecher,  1886)' 

Amnicola  antroeretes  (Hubricht,  1940)' 

Amnicola  corae  Hubricht,  1979' 

Am.nicula  prosperpiria  Hubricht.  194(1' 

Amnicola  stygia  Hubricht,  197P 

Antrolna  culvert  Hubricht,  197 P 


'Pyrgulopsis  ozarkertsis  is  placed  in  Ma  rston  ia  following  the 
suggestion  of  Thompson  (1977). 

'Burch  and  Tottenham  (1980)  included  Amnicola  mi.s.sowrj- 
ensis  Pilsbry,  1898  as  a  valid  species.  Although  this  group  re- 
quires further  anatomical  investigation,  the  argument  of 
Hubricht  (1940)  for  synonymizing  this  species  under  .4. 
aldrichi  is  followed. 
^Troglobitic. 


relationships  to  the  hydrobiid  subfamilies 
possessing  single-duct  penes  (e.g.  Lithoglyphi- 
nae).  Additionally,  Thompson  (1984)  suggested 
that  the  simple,  non-appendaged  penis  was  a 
generalized,  primitive  condition  in  the 
Lithoglyphinae  but  may  occur  as  a  "derived  con- 
dition through  the  secondary  loss  of  previously 
existing  characters"  for  some  taxa  in  other 
hydrobiid  subfamilies  (e.g.  Hershler  and  Davis, 
1980:  Hydrobiinae;  Hershler,  1985:  Littoridini- 
nae).  Considering  similarities  of  the  antrobian 
radula,  shell,  and  operculum  to  Amnicola. 
morphology  of  the  penis  may  represent  a 
secondarily  derived  character  state  mAntmbid. 
In  conjunction  with  the  above,  a  conservative 
classification  would  retain  A.  culveri  in  the 
Amnicolinae  until  phylogenetic  relationships 
can  be  resolved. 

The  only  endemic,  epigean  aninicoline  pre- 
sently recognized  \?,  Amnicola  aldrichi  (Call  and 
Beecher),  which  exhibits  a  habitat  preference 
for  springs.  Hubricht  (1940)  described  three 
subsj)ecies  ior  A.  aldrichi:  aldrichi  s.s.,  itisolUa 
Hubricht,  and  antroec.ete.a  Hubricht.  "Deeper 
sutures,  strongly  shouldered  whorls,  and  a  free 


lip  (Hubricht,  1940)"  differentiated  insolita  from 
aldrichi  s.s.  Hubricht's  illustration  and  descrip- 
tion of  a  free  lip  indicate  a  slightly  uncoiled  body 
whorl.  This  condition  occurs  occasionally  within 
the  Gastropoda  and  has  been  shown  to  some- 
times represent  distinct,  geographical  sub- 
species (e.g.  Clarke,  1973:  Valvata  sincera  on- 
tariensis  Baker).  Recently  collected  specimens 
(juveniles-adults)  from  one  of  Hubricht's  insolita 
localities  (2  November,  1985;  Meramec  Spring, 
8  km  southeast  of  St.  James,  Phelps  County, 
Missouri)  were  examined  and  found  to  exhibit 
considerable  variability  with  regards  to  the  dis- 
tinguishing characters  for  insolita.  In  no  case 
was  there  any  evidence  of  an  uncoiled  body 
whorl.  Palmer  (1985  and  references  within) 
demonstrated  that  shell  variation  in  gastropods 
can  be  strongly  influenced  by  environmental 
conditions  and/or  intraspecific  genetic  plasitici- 
ty  and  does  not  necessarily  reflect  divergence 
(i.e.  speciation).  The  distribution  of  insolita 
(from  Hubricht,  1940)  is  not  geographically 
distinct  from  that  of  aldrichi  s.s.  Considering 
the  above,  itisolifa  does  not  appear  to  warrant 
subspecific  status  in  the  modern  sense  (see 
Mayr,  1966). 

Morton  (1967)  and  many  others  have  dis- 
cussed problems  related  to  constructing  mollus- 
can  classifications  from  single-based  systems 
(e.g.  shell  morphology).  As  noted  above,  varia- 
tion in  shell  characters  does  not  always  imply 
speciation  (Palmer,  1985).  Conversely,  lack  of 
shell  morphological  variance  is  not  always  in- 
dicative of  conspecificity  or  close  phylogenetic 
relationships,  but  may  mask  divergent  or  con- 
vergent evolution  (Davis,  1979;  Dillon  and 
Davis,  1980;  Vail,  1980;  Thompson,  1984). 
Hubricht  (1940)  was  unable  to  readily  differen- 
tiate shells  of  his  third  subspecies,  Amnicola 
aldrichi  antroecetes.  from  slender  examples  of 
aldrichi  s.s.  Both  the  previously  discussed 
"subspecies"  are  epigean  forms  with  pigmented 
bodies  and  functional  eyes;  however,  antroecetes 
is  an  unpigmented,  blind,  obligate  troglobite. 
Hubricht  (1940)  failed  to  be  consistant  in  his 
methodology  for  distingTiishing  the  subspecies 
of  A.  aldrichi:  insolita  i)y  shell  characters  vs. 
antroecetes  by  anatomy  and  operculum.  Even 
though  insolita  is  considered  invalid,  this  devia- 
tion still  suggests  that  different  levels  of 
organization  were  being  examined.  Thompson 


100(1 


At)iMl  :-!(),  Ut8C. 


THE  NAUTILUS 


it 


(1984)  indicated  that  "a  hi.u'h  degree  of 
anatiitnical  uniformity"  exists  within  groups  of 
the  Hydrobiidae  (e.g.  Lithoglyi.)hinae).  The  loss 
of  pigmentation  and  eyes  represents  a  rather 
radical  divergence  from  the  epigean  anatomy  of 
aldrichi  s.s.  To  consider  these  troglohitic  ada]> 
tations  to  be  ecophenotypic  variation  implies  an 
amazing  genetic  plasticity  and  an  "use  it  or  lose 
it"  evolutionary  interpretation.  Ecophenotypic 
variation  does  not  meet  modern  criteria  for  sub- 
specific  status  (Mayr,  1966);  however,  previous 
relegation  of  nntroeretes  as  a  subspecies  reflects 
the  over-reliance  on  shell  characters  for  con- 
structing phylogenies.  Amnicola  antroecefes 
should  be  considered  a  distinct  species.  In- 
terestingly, A.  iintriieceteii  has  a  rather  wide 
distribution  when  compared  to  other  regional 
troglobitic  gastropods  (restricted  to  single  or 
connected  caves).  Other  troglobitic  taxa  (e.g. 
Amblyopsidae;  Isopoda:  Caecidotea)  exhibit 
similarly  wide  ranges  in  the  Interior  Highlands. 
This  may  be  related  to  the  karst  geology  of  the 
region.  Hubricht  (1940)  did  note  some  size  varia- 
tion at  different  localities  liut  correlated  this  to 
food  availability.  However,  due  to  its  wide  range 
he  suggested  that  .4.  antn)ecefe><  may  represent 
a  composite  group.  Further  investigation  is  ob- 
viously warranted  for  the  A.  aldrichi  complex 
and,  for  that  matter,  the  Hydrobiidae  through- 
out the  Interior  Highlands. 

Acknowledgments 

The  uniqueness  of  Somatogyrus  roi^ewnteri 
was  confirmed  by  Fred  G.  Thompson,  Florida 
State  Museum,  University  of  Florida.  SEM's 
were  prepared  with  the  assistance  of  Claudia  S. 
Bailey  and  specimens  o{ Amnicola  aldrichi  were 
provided  by  Michael  L.  Mathis,  both  Depart- 
ment of  Zoology,  University  of  Arkansas. 


LITERATURE  CITED 
Baker.  F.  C.  1928,  The  fresh-water  Mollusca  of  Wisconsin, 

part  1:  Gastropoda.  BulL   Wiitc.  Geol.  Nat.  Hist.  Sun: 

70:1-507. 
Burch.  J.  B.  and  J.  L.  Tottenham.  1980.  North  American 


Ireshvvatcr  snails;  species  Hst.  runnes,  and  illustratinns. 
W.ilbrniir,  \:S]--l\r,. 

Clarke,  A.  11.  197M.  The  freshwatei'  molluscs  of  the  Cana<lian 
Interior  Basin.  Mulnaihiyin  13:1-509. 

Davis,  G.  M.  1979.  The  origin  and  evolution  of  the  gastropod 
family  Pomatiopsidae,  with  emphasis  on  the  Mekong  River 
Triculinae.  Aaid.  Nut.  Sri.  Phil.  Monoiji:  20:1-120. 

Dillon.  R.  T.  and  G.  M,  Davis,  1980.  The  Gimwlmxiit  of  south- 
ern Virginia  and  northwestern  North  (Carolina:  genetic 
and  shell  morphometric  relationshijis.  Mnlacolitijtti 
20:83-98. 

Gordon,  M.  E.  1980,  p'reshwater  Mollusca  of  the  Elk  River, 
White  River  above  Beaver  Reservoir,  and  Frog  Bayou 
drainages  of  the  southwestern  Ozarks.  Unpublished  M,S, 
thesis.  University  of  Arkansas,  .366  pp. 

1981,  Recent  Mollusca  of  Arkansas  with  annota- 
tions to  systematics  and  zoogeography,  Proc.  Ark.  Anid. 
Sri.  34(1980):.5S-62. 

Hinkley,  A,  A,  1915,  New  fresh-water  shells  from  the  Ozark 
.Mountains.  Pruc.  U.S.  Nail.  Mm.s,  49:587-589. 

Hershler.  R,  1985,  Systematic  revision  of  the  Hydrobiidae 
(Gastropoda:  Rissoacea)  of  the  Cuarto  Cienegas  basin, 
Goahuila.  Mexico,  Malacologia  26:31-123, 

Hershler,  R,  and  G.  M.  Davis,  1980,  The  morphologv'  of 
Hudrobia  Irunrata  (Gastropoda:  Hydroliiidae):  relevance 
to  systematics  of  Hi/drohia.  Biol.  Bull.  158:19.5-219. 

Huliricht,  L.  1940,  The  Ozark  amnicolas.  Tin-  Naiililiis 
53:118-122, 

1971,  New  Hydrobiidae  from  Ozark  caves,  77/r 

Naiitilii.<:  83:93-96, 

1979,  A  new  species  of  Amnicola  from  an  Arkan- 


sas cave  (Hydrobiidae).  The  Naalil ua  94:14S 
Mayr.  E.  1966.  Animal  species  and  evolutinn.  The  Belknap 

Press  of  Harvard  University  Press,  Cambridge.  797  \)]). 
Morton,  .J,  E,  1967,  Molluscs.  Hutchinson  and  Company, 

Ltd,,  London,  244  pp. 
Palmer,  A,  R,  1985,  Quantum  changes  in  gastropod  shell 

morphology     need     not     reflect     speciation,     Eroliitiou 

39:699-705, 
Sampson.  F,  A,  1913,  A  preliminary  list  of  the  Mollusca  of 

Missouri  (exclusive  of  the  Unioniade),  Tmn.s,  Acad.  Sci. 

St.  Lo)(),s  22:67-108, 
Thompson.   F,  G,    1977,  The  hydrobiid  genus  Marstonia. 

Bull.  Fla.  Slate  Mus.  (Bi(j|,  Sci,)  21:113-1.58, 
1984,  North  American  freshwater  snail  genera 

of  the  hydrobiid  subfamily  Lithoglyphinae.  Malacologia 

25:109-141, 
Vail.  V,  A,  1980.  The  species  problem  in  Campeloiiia  (Gas- 
tropoda:    Viviparidae),     Bull.    Amer.     Malacol.     Union 

1979:67, 
Walker,  B,   1915,  Apical  characters  in  Somatogyrus.  with 

descriptions  of  three  new  species.  The  Nautilus  29:37-41, 

49-.53, 
Wheeler,  H,  E,  1918.  The  Mollusca  of  Clark  County.  Arkan- 
sas. The  Nautilus  31:109-125. 


78      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


FAVARTIA  BREVICULA  (SOWERBY,   1834)  AND  TWO  NEW  SPECIES 

OF  FAVARTIA  FROM  THE  WESTERN  PACIFIC 

(GASTROPODA:  MURICIDAE) 


Anthony  D'Attilio  and  Barbara  W.  Myers 

Department  of  Marine  Invertebrates 

San  Diego  Natural  History  Museum 

San  Diego,  CA  92112 

ABSTRACT 
Murex  breviculus  Sowerby,  18SJt,  is  reexamined  and  a  lectotype  is  selected  from 
the  syntypic  lot.  Murex  tetragonus  Broderip,  1833,  is  discussed  and  it  is  our  con- 
clusioyi  the  name  is  a  nomen  dubium.  Two  7iew  species  of  Favartia  Jousseaume, 
1880,  are  described,  one  from  the  Solomons  one  from  the  Philippines. 


The  identity  of  Murex  breviculv^s  Sowerby, 
1834,  has  never  been  in  doubt  and  the  figure  in 
the  Conrhological  Illustrations  (Sowerby,  1834, 
pi.  63,  fig.  37)  clearly  defines  the  species.  The 
illustration  agrees  with  the  brief  but  adequate 
description  published  later  in  the  Proceedings  of 
the  Zoological  Society  of  London  (1841:146-i47J. 
The  syntypic  lot  consisting  of  three  specimens 
was  borrowed  from  the  British  Museum.  A 
lectotype  was  selected  and  the  two  remaining 
specimens  are  considered  paralectotypes.  For 
purposes  of  clarity  all  three  specimens  from  the 
type  lot  are  illustrated  herein  through  the 
courtesy  of  the  Trustees  of  the  British  Museum 
(Natural  History). 

The  identity  of  Murex  tetragonus  Broderip, 
1833,  on  the  other  hand  has  never  been  estab- 
lished. Broderip's  description  in  the  Proceedings 
of  the  Zoological  Society  of  London  for  1832, 
was  based  on  a  single  "very  much  waterworn" 
specimen.  Broderip  did  not  figure  this  species, 
but  Sowerby  (1834)  figured  M.  tetragonus  in  the 
Conchological  Illustrations  (pi.  61,  fig.  25  and  pi. 
63,  fig.  36).  We  were  informed  by  Ms.  Kathie 
Way  of  the  Molluscan  Section,  British  Museum, 
that  they  were  unable  to  locate  any  type 
material  of  M.  tetragonus  Broderip. 

Institutional  abbreviations  used  in  this  paper 
are:  ANSP  =  Academy  of  Natural  Sciences 
Philadelphia.  BM(NH)  =  British  Museum 
Natural  History.  SDNHM  =  San  Diego  Natural 
History  Museum. 

Muricidae  Rafinescjue,  1815 
Muricopsinae  Radwin  and  D'Attilio,  1971 


Favartia  Jousseaume,  1880 

Type  Species:  Funirtin  brcricula  (Sowerby,  1834) 
by  original  designation 

Favartia  brevicula  (Sowerby,  1834) 

Murex  brevirulus  Sowerby,  1834:  Conch.  111.  PI.  63.  fig.  37 
Figs.  1-8,  13 

The  syntypic  lot  oi Favartia  brevicula  (Sower- 
by, 1834)  borrowed  from  the  British  Museum 
(NH)  consisted  of  three  beach  eroded  specimens 
reg.  #1974088. 

The  lectotype,  measuring  26  mm  x  20  mm, 
was  selected  because  it  had  the  least  amount  of 
erosion.  Shell  is  broadly  biconic,  the  number  of 
whorls  indeterminate  because  of  the  erosion  of 
the  spire.  Whorls  convex,  aperture  subcircular, 
smooth  within,  inner  lip  erect,  outer  lip  crenu- 
late,  reflecting  external  cords,  anal  sulcus  not 
apparent,  canal  narrowly  open,  broad,  moderate 
in  length,  terminally  sharply  recurved:  three 
older  canal  terminations  on  fasciole;  four  broad 
varices  crossing  shoulder  and  abutting  whorl 
above;  narrow  intervarical  areas  appear  smooth 
on  lectotype;  leading  edge  of  varices  reaching 
midway  in  the  intervarical  area;  varices  deeply 
cut  and  depressed  on  both  leading  and  receding 
side;  where  varix  crosses  the  shoulder  it  is 
bladelike  and  strongly  recurved;  the  leading  side 
of  the  varix  with  longitudinal  lamellae  un- 
dulating between  the  spiral  cords;  five  thick 
spiral  cords  on  body  whorl,  obsolete  in  narrow 
intervarical  areas,  the  first  three  about  equal 
size,  the  two  anterior  ones  small  and  grouped  as 
a  pair.  Two  minor  cords  on  the  canal  separated 
by  a  gap  from  the  body  cords  at  the  base  of  the 
l)ody  whorl.  Color  of  types  is  dull-white. 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      79 


p'KiS.  l-(i.  Fiirnrtiii  hriTiruUi  (Siiwerliy.  1X34). 
1,  Dorsal  and  2,  apertural  views  of  lectotype 
BM(NH)  #lii74088  (26.0  x  20.0  mm.);  3  and  4, 
paralectotype  BM(NH)  #1974088  (23.75  x  17.0 
mm.);  5  and  6.  paralectotype  BM(NH)  #1974088 
(26.75  X  20.0  mm.). 


FIGS.  7  and  8.  Farartlit  hirrirulii  (Sowerby.  1834) 
SDNHM  76669  small  specimen  from  Bohol  Straits,  Philip- 
pine Islands.  7,  protoconch;  8,  general  shell  morphology 
(11.0  mm  X  7.4  mm.). 


Type  Locality -Unknown.  The  species  if 
found  throughout  the  western  and  south  Pacific 
to  east  Africa. 


Dimensions  - 
BM(NH)  #1974088  ■ 
BM(NH)  #1974088 
BM(NH)  #1974088  ■ 


Length     Width 

26.0  mm  X  20.0  mm 

23.75  mnix  17.0  mm 

26.75  mm  X  20. 0  mm 


Lectotype 

Paralectotype 

Paralectotype 

DisciLSsion - So-werhy  (1879)  in  the  Thesaurus 
Conchyliorwn  appeared  to  change  the  descrip- 
tion of  F.  hrevicula.  Instead  of  having  four 
varices  as  described  in  the  original  description, 
he  stated  there  were  five  varices  for  F. 
hrevicula.  Although  the  three  syntypes  for  F. 
brevicula  have  only  four  varices  and  in  most  of 
the  specimens  examined  there  were  only  four 
varices,  among  the  material  at  our  disposal,  we 
found  nine  specimens  with  five  varices  from  dif- 


19  :-  ■,  20 

FIGS.  19  and  20.  F.  Mmgonn  Broderip.  1833,  taken  from 
Sowerby  (1834).  19,  from  pi.  61,  fig.  25;  20,  from  pi.  63, 
fig.  36. 


80      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


ferent  localities  and  ranging  in  size  from  11.5 
mm  to  23.0  mm.  The  number  of  varices  appears 
to  be  a  variable  character. 

Favartia  tetragona  (Broderip,  1833) 

Murex  tetragonus  Broderip.  1833:  Proc.  Ciimm.  Sci. 

Corresp.  Zool.  Soc.  London.  174. 

Figs.  19,  20 

The  single  specimen  available  to  Broderip  was 

in  poor  condition  and  the  only  one  he  ever  saw. 

It   was   never   figured   by    Broderip   and   the 

description    is    not    definitive.    The    original 

description  is  as  follows: 

Mliiie\    TETiiAUONl'l,.      MuT .   tcstd  pt/Tamtdali,    albidA,   ouadri- 
Jaruim   tancosn,    tranivrrtim    caslatu,    suLcaucellatu ;    apcrturit 
violacca,  protmncnte  ;    iabii  Umbo  unduloio-crmutato  ;    canali 
Lrevi,  recurvd ;   long.  \i,  lot.  J  poll. 
Hab.y 

Mu£   Sowerbv 
'J'his   sppcinicn.   the  onlv   one  1    ever  saw,  is  ver\'  much   watet- 

worn,  but  llie  leadinfr  cliarnclers  of  lliu  tuecie.',  ore   uiiiniurtL  

W.  J.  I.. 


According  to  Broderip  the  habitat  was  un- 
known. The  specimen  was  deposited  in  Museum 
Sowerby  and  its  whereabouts  today  is  unknown. 
The  illustrations  by  Sowerby  in  1834  (PI.  63,  fig. 
36  and  PL  61,  fig.  25)  do  not  show  a  waterworn 
shell  as  described  by  Broderip,  but  do  show  a 
very  pyramidal-shaped  shell,  with  a  broad  base 
and  flaring  lip.  Figs.  19  and  20  are  copied  from 
Sowerby  (1834). 

Lacking  typological  material  the  morphology 
of  this  taxon  must  rely  entirely  on  Broderip's 
description  and  the  figure  by  Sowerby.  Reeve 
(1845:  PI.  26,  fig.  18)  figured  the  broad  coarse 
shell  of  Murex  breviculuH  Sowerby  placing  M. 
hreviculus  in  the  synonymy  of  M.  tetragonus.  the 
earlier  name,  stating  he  could  not  discover  "any 
s{)ecific  differences  between  Murex  tetragonuH 
and  hreviculuH".  Sowerby  who  did  the  illustra- 
tions for  the  Conchologia  Iconica  did  not  dispute 
the  synonymy.  However,  in  1879,  in  the 
Thesaurus  Conchyliorum.  Sowerby  included 
both  binomens  {Murex  tetragonuti  sp.  187,  fig. 
228  and  M.  hreviculus  sp.  188,  fig.  229),  and  in 
comparing  the  two  species  stated  at  that  time 
that  M.  hreviculus  was  a  shorter  and  thicker 
shell  than  M.  tetragonus  and  had  five  varices. 
This  differed  from  M.  tetragonus  which  was  de- 
scribed as  having  four  varices.  In  1879  Sowerby 
also  stated  for  M.  tetragonus  that  the  latticed 
varices  follow  from  whorl  to  whorl  so  conflu- 


ently  as  to  form  four  oblique  walls  up  the  spire 
and  he  stated  for  M.  breviculu^s  "ad  spiram  inter- 
ruptis". 

The  identification  of  M.  tetragonus  has  never 
been  established.  As  stated  above  Reeve  (1845) 
synonymized  M.  hreviculus  with  M.  tetragonua. 
the  earlier  name.  Tryon  (1880:  pi.  36,  fig.  424, 
425)  agreed  with  Reeve,  figuring  both  the  broad 
heavy  shell  of  Af.  hreviculus  and  what  looks  like 
a  copy  of  Sowerby's  figure  of  M.  tetragonus, 
both  under  the  name  of  M.  tetragonus. 

In  recent  works  the  following  authors  figured 
what  they  believed  to  beF.  tetrngona  as  follows: 

Smith,  M.  (1939:  pi.  6,  fig.  2)  figured  F. 
hrevicula. 

Cernohorsky  (1967:  pi.  15,  fig.  20)  figured  an 
undetermined  species. 

Fair  (1976:  pi.  20,  fig.  295)  figured  an  undeter- 
mined species. 

Radwin  and  D'Attilio  (1976)  mistakenly  de- 
scribed F.  tetrngona  as  having  three  varices  and 
figured  it  on  pi.  14,  fig.  3.  This  is  the  new  species 
described  here  as  F.  trivaricosa  and  shown  also 
in  text  fig.  96  of  Radwin  and  D'Attilio. 

Inasmuch  as  there  is  apparently  no  type  ex- 
tant for  F.  tetragona,  the  identity  of  the  species 
has  not  been  established  in  150  years.  We  have 
not  been  able  to  associate  Sowerby's  figure  with 
any  species  of  Favartia  known  to  us  and  we 
believe  this  species  is  of  dubious  merit,  therefore 
we  are  placing  it  among  the  doubtful  names.  It 
is  interesting  to  note  that  no  authors,  to  date, 
have  mentioned  Broderip's  description  of  the 
aperture  being  prominently  violet. 

Favartia  robertsoni  new  species 

Figs.  9-12,  14,  15.  16 

Shell  medium  size,  broadly  fusoid,  spire  mod- 
erately high,  anterior  truncate,  approximately 
five  whorls,  early  whorls  eroded;  protoconch  not 
determined;  weakly  shouldered,  somewhat  tabu- 
late; shoulder  area  between  the  varices  de- 
pressed and  concave,  especially  on  receding  side 
of  the  varix;  suture  impressed;  body  whorl 
broader  than  high;  aperture  subovate;  peris- 
tome erect;  outer  lip  undulate,  reflecting  exter- 
nal transverse  cords;  canal  broad,  very  narrowly 
open,  short,  terminally  tube-like,  bent  backward 
nearly  at  right  angle;  siphonal  fasciole  retaining 
two  older  canal  terminations. 

Varices   thick,    four   per   whorl;    intervarical 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      81 


■1 

1 

1 

^^^^^^^MiS^ 

w'PV 

^^1 

W^^ 

i^ 

\% 

HP^^Hk-^^ 

^"^^ 

-^i^k 

Ei^^        V^^jj|i 

mm 

■ 

Kf^*^ 

mm 

H 

^^^1^^^^^ 

■ 

1 11 1 

FIGS.  9-12.  9,  10,  Fiivartin  rohi-ii^oiu  n.  sp.  holotype  SDNHM  86944  (26.0  > 
26.5  mm.).  11  and  12,  paratype  Din  the  Donald  Pisor  collection  (26.6x27.2  mm.; 


areas  with  moderately  raised  cords;  varices  ex- 
tending over  shoulder  obliquely  to  left  and  ter- 
minating at  suture. 

Spiral  sculpture  as  follows:  one  broad  cord  at 
the  shoulder  prolonged  prominently  into  a  blunt 
spine,  the  margin  nearly  closed  on  the  leading 
side;  a  similar  broad  cord  below  extending 
beyond  the  varix,  but  not  prolonged  into  a  spine, 
up  to  the  penultimate  varix.  On  the  mature 
apertural  varix  these  two  cords  do  not  project 
beyond  the  remaining  sculpture  of  the  varix. 
Four  secondary  cords  on  the  body  whorl  dimin- 
ishing in  size  anteriorly;  varix  deeply  excavated 
between  the  cords  on  receding  side;  cords  on 
body  whorl  more  or  less  weakly  raised;  three 
minor  cords  on  shoulder  obsolete  except  near 


apertural  varix.  Leading  side  of  varices  re- 
curved and  with  scabrous  lamellae  producing  a 
surface  with  squarish  pits;  weak  scabrous 
lamellae  on  surface  of  spiral  cords  between 
varices. 

Color:  Creamy  white  entirely  except  for  the 
holotype  with  is  a  dead  collected  shell  lightly 
stained  with  rust. 

Type  Locality -Ma.\si\t2.  Island,  Solomon 
Islands. 

Dimensions-  LenKth    width 

Holotype  SDNHM  8B944 

.Ataa,  Malaita  Island,  Solomon  Is.  2t;.l)         21)  5  mm 

Paratype  A,  SDNHM  86942 

Marau  Sound,  Guadalcanal,  Solomon  Is.  17,0         1,5,',) 

I'aratype  B,  ANSP  289.'JBK 

Malaita  Island,  Solomon  Islands  2(1, .5         20,2 


82      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


KKjS.  lIMti.  13,  Outline  drawinn'  <if  Furnrtiii  hrcriruhi 
(Sowerby,  1884)  SDNHM  78704  (19.6  x  15.2  mm.).  FIGS. 
14-16,  Fnvartia  nibcrtsoni  D'Attilio  and  Myers.  14, 
paratype  B  ANSP  289568  (20.5  x  20.2  mm.);  15,  paratype  C 
ANSI'  289568  showing  micro.sculpture  of  receding  side  of 
varix;  16,  leading  side  of  varix. 


f'aratyiif  C,  ANSI'  '289568 

Malaita  Island.  Solomon  Islands  14.4        14  1 

Paratype  D.  Donald  Pisor  collection. 

Malaita  Island.  Solomon  Lslands  26.6         27.2 

Paratype  E,  Glass  and  Foster  collection 

Marau  Sound.  t'lUadalcanal,  .Solomon  is.  24.5         1.75 

Etymology -This  species  is  named  for  Dr. 
Robert  Robertson,  Curator  and  holder  of  the 
Pilsbry  Chair  of  Malacology',  Academy  of 
Natural  Sciences  Philadelphia,  whose  mala- 
cological  insight  the  senior  author  has  long  ad- 
mired. 

D i.sTM.s.s?o«  -  The  varices  appear  the  same  ex- 
cept when  fully  mature;  it  is  only  at  maturity 
that  the  final  varix  shows  its  character  as  in  the 
above  description.  Comparison  is  made  with  its 
nearest  congener,  F.  brevicula  (Sowerby,  1834), 
which  has  a  thicker,  coarser  shell  with  a  broader 
more  obese  body  whorl  and  larger  aperture.  F. 
brevicula  lacks  the  extended  long  heavy  spine  on 
the  varices  previous  to  the  mature  apertural 
varix.  It  also  has  no  noticeable  keel  on  the 
shoulder  and  has  five  equal  cords  on  the  body 
whorl  in  contrast  to  the  two  major  cords  and 
four  secondary  cords  of  F.  robertson  i. 

F.  robertsoni  new  species  was  figured  in 
Radwin  and  D'Attilio  (1976)  pi.  23,  fig.  1 1  and  pi. 
24,  fig.  2  as  F.  brevicula  (Sowerliy,  1834). 

Favartia  trivaricosa  new  species 

Figs.  17,  18 

Shell  small  fusoid,  spire  high,  anterior  trun- 
cate, approximately  seven  whorls:  protoconch 
eroded,  not  determined;  suture  impressed. 
Aperture  ovate,  peristome  erect  anteriorly,  but 
appressed  posteriorly  on  the  inner  side;  outer  lip 
crenulate  reflecting  external  transverse  cords, 
canal  short  very  narrowly  open,  terminally  tube- 
like and  bent  backward  nearly  at  right  angle, 
siphonal  fascicle  retaining  one  older  canal  ter- 
mination. 

Varices  moderately  thick,  three  per  whorl  on 
the  final  and  penultimate  whorl,  four  on  the 
earlier  whorls;  varices  extending  over  shoulder 
and  terminating  at  suture.  Intervarical  areas 
with  axial  sculpture  in  the  form  of  low,  flat, 
thickened  lamellae.  Sjiiral  sculpture  as  follows: 
one  cord  above  the  shoulder  and  four  broad 
cords  on  the  Ixxly  whorl.  Leading  side  of  the 
varices  recurved  and  with  scal)rous  lamellae, 
deeply  excavated  between  the  cords  on  the 
receding    side.    A    combination    of   the    axial 


Vol.  100(2) 


April  30,  1986 


THE  NAUTILUS      83 


FICS.  17  and  18.  Famrtid  trivancoxa  n.  sp.  hulotype  SDNHM  86943  (15.0  x 
VI. 4  mm.). 


lamellae  crossed  by  these  spiral  cords  results  in 
square  pits  in  the  intervarical  areas.  When  worn 
only  the  pits  are  visible. 

Color:  Dull-white  varices  with  very  pale-tan 
intervarical  areas;  aperture  pale  lavender. 

Type  Locality-Boho\  Straits,  Philippine 
Islands. 


Drmensions- 

Leiiifth 

Width 

nini. 

mm. 

Hcilotype  SDNHM  8ill»43 

Bohol  Straits.  P.  1.  (ex  Foster  and  Class) 

\bM    X 

0.4 

Paratype  A,  SDNHM  8205)4 

Celm  Island,  P.  1. 

13..5  X 

9.4 

Paratype  B,  SDNHM  737(12 

Bohol  Straits.  P.  I. 

13.0  X 

H..'') 

Paratype  C.  SDNHM  76(i(i8 

Mactan  Island,  P.  I. 

12.0  X 

fi.B 

Paratype  D.  SDNHM  8227e 

Mactan  Island.  P.  I- 

14.3  X 

9.7 

Paratype  E. 

Glass  and  Foster  eollection.  P.  I. 

13.0  X 

8.7 

Other  Material  Era  mined -One  specimen  in 
the  June  King  collection  from  Bohol  Straits,  P. 
I.  (11.0  X  6.0  mm.);  one  specimen  in  the  Rose 
D'Attilio  collection  from  Wading  Island,  Fiji 
Islands  (18.3  x  11.4  mm). 

Discussion  -  Comparison  is  made  with  the  fol- 
lowing species  ol  Favartia: 

F.  alveata  (Kiener,  1842)  from  the  West  Indies 
which  has  five  to  seven  varices  instead  of  three 
as  does  F.  trivaricosa,  and  spiral  cords  orna- 
mented with  scales  appressed  to  their  surface. 

F.  brevicula  (Sowerby,  1834)  which  is  a  broad 
heavy  larger  shell  with  four  or  five  broad  sculp- 
tured varices  and  smooth  intervarical  areas. 


F.  cyclostoma  (Sowerby,  1841)  which  is  a  large 
shell  with  five  varices. 

F.  minatauros  Radwin  and  D'Attilio,  1976 
from  Mauritius  which  has  an  obtuse  spire  and  a 
swollen  body  whorl. 

F.  mundit  (Reeve,  1849)  from  Japan  which  has 
four  to  five  varices. 

F.  trivaricosa  new  species  was  mistakenly 
figured  in  Radwin  and  D'Attilio  (1976)  pi.  14, 
fig.  3  and  text  fig.  96  as  F.  tetragona  (Broderip, 
1833). 

Etymology -Th\s  new  species  is  named  for  the 
three  varices  on  the  body  whorl  which  distin- 
guish it  from  several  of  its  congeners.  Latin  tri 
(three)  plus  varicosa  (varices). 

Acknowledgments 

We  wish  to  thank  the  following  for  the  loan  of 
typological  and  comparison  material:  Academy 
of  Natural  Sciences  Philadelphia;  American 
Museum  of  Natural  History,  British  Museum 
(Natural  History);  Rose  D'Attilio;  June  King  and 
Donald  Pisor  of  San  Diego,  California;  and 
Charles  Glass  and  Robert  Foster  of  Santa 
Barbara,  California.  We  also  thank  David  K. 
Mulliner  for  the  photography  used  in  this  paper. 
William  K.  Emerson  and  Walter  E.  Sage,  III 
kindly  reviewed  the  paper. 

LITERATURE  CITED 

Broderip,  W.  ,1.  183.3.  Characters  of  new  species  of  Mollusca 
and  Conchifera  collected  liy  H.  Cuming.  Proc.  Cunim.  Sci. 
Corrpitp.  Ziiiil.  S'ic.  Lmuldii  for  1832.  pp.  173-179. 


84      THE  NAUTILUS 


April  30,  1986 


Vol.  100(2) 


Cernohorsky.  W.  ().   1967.  The  Muricidae  of  Fiji.  Vi'ligcr 

10(2):1 11-132,  pis.  14-15. 
Fair,  R.  1976.  The  Murcx  Book:  an  illustrated  catalogue  (if 

Recent  Muricidae  (Muricinae.  Muricopsinae.  Ocenelirinae) 

138  pp.,  23  pis. 
.lousseaume,  F.  1880.  Division  methodiqiie  de  la  famille  des 

Purpurides.  Le  Naturaliste  42:335-336. 
Kiener,  L.  C.  1842-43.  Species  general  et  icdnoyniphie  ilea 

ciKjiiiUen  ririniti's  7,  Rocher  (MH/v':r),  pp.  1-130,  pis.  1-47. 
Fiadwin.   G.   and   A.   U'Attilio.    1976.  Murex  shells  of  the 

World,  an  illustrated  guide  to  the  Muricidae.  Stanford 

University  Press.  284  pp..  32  pis.,  192  text  figs. 
Rafinesque,  C.  S.  1815.  Analyse  de  la.  nature  ou  tableau  dii 

univers  et  des  corps  organises.  Barravecchia,  Palermo. 


Reeve,  L.  1849.  Conch.  Iconica  vol.  Ill,  Murex,  errata  ad 

fiiirni.  Index. 
Smith,  M.  1939.  An  illustrated  cataloij  of  the  recent  species 

of  the  rock  shell.  Muricidae,  Thaisidae  and  Coralliophilidae. 

Lantana,  Florida.  34  pp.,  21  pis. 
Sowerby,  G.  B.  (second  of  name).  1834.  Conch.  Illust.  Murex. 

London,  pis.  .58-67. 
1841.    Descriptions   of  some   new   species  of 

Murex.  principally  from  the  collection  of  H.  Cuming,  ESQ. 

Pj'oc  ZooI.  Soc.  London,  pp.  137-147. 
1879.  Thesaurus  Conrhyliorum.  London,  vol.  4, 


55  pp.,  24  pis. 

Tryon,  G.  W.,  .Ir.  1880.  Manual  ofConcholoipi  Philadelphia, 
vol.  2,  289  pp.,  70  pis. 


ANTIQUARIAAT  JUNK  B.V. 

Van  Eejfhenstraat  129 

1071  GA  AMSTERDAM 

The  Netherlands 


NATURAL  HISTORY  BOOKSELLER  SINCE   1899. 

.lust  issued  a  si)ecial  list  of  RARE  and  VALUABLE  early  books  on  SHELLS. 

OtTered  among  others  are:  BORN  -  Testacea  Musei  Caesarei  Vindohonensis. 
Vienna  1780;  BROWN  -  Illustrations  of  the  Conchology  of  Great  Britain. 
London  1827;  BUONANNI  -  Recreatione  dell  Occhio  .  .  .  Rome  1681; 
PERRY  -  Conchology.  London  1811;  REEVE  -  Conchologia  Iconica  .  .  .  185 
parts  (of  289).  London  1843;  SOWERBY  -  Mineral  Conchology,  and  many 
other  fine  handcoloured  illustrated  works.  Please  write  if  you  wish  to  receive 
our  special  list. 


INFORMATION  FOR  SUBSCRIBERS 


The  annual  subscription  rate  for  The  Nautilus 
is  $15.00  for  individuals  (foreign  $18.00)  and 
$20.00  for  institutions  (domestic  or  foreign). 
Subscriptions  may  begin  in  January.  Send  check 
or  money  order  made  out  to  "American  Mala- 
cologists"  to  the  Business  Manager,  P.O.  Box 
2255,  Melbourne,  Florida  32902-2255,  U.S.A. 


Back  issv£s  from  volume  72  to  date  are  ob- 
tainable from  the  Business  Manager.  Volumes  1 
through  71  (if  available)  may  be  obtained  in 
reprint  or  original  form  from  Kraus  Reprint 
Co.,  Route  100,  Millwood,  New  York  10546. 

Advertising  rates  may  be  obtained  from  the 
Business  Manager  or  Editor. 


CONTRIBUTORS 


Manuscripts:  Authors  are  requested  to  follow 
the  recommendations  of  the  Style  Manual  for 
Biological  Journals,  which  may  be  purchased 
from  the  American  Institute  of  Biological  Sci- 
ences, 1401  Wilson  Boulevard,  Arlington,  Va. 
22209.  Manuscripts  should  be  typewritten  and 
doublespaced;  original  and  one  copy  are  re- 
quired, to  facilitate  reviews.  Tables,  numbered 
in  arable,  should  be  on  separate  pages,  with  the 
title  at  the  top.  Legends  to  photographs  should 
be  typed  on  separate  sheets.  Explanatory  terms 
and  symbols  within  a  drawing  should  be  neatly 
printed,  or  they  may  be  pencilled  in  on  a  translu- 


cent overlay,  so  that  the  printer  may  set  them  in 
8  pt.  type.  There  is  a  charge  of  50  cents  per 
word  for  this  extra  service.  All  authors  or  their 
institutions  will  be  charged  50  cents  per  line  of 
tabular  material  and  taxonomic  keys.  The  pub- 
lishers reserve  the  right,  seldom  exercised,  to 
charge  $45  per  printed  page. 

An  abstract  should  accompany  each  paper. 

Reprints  are  available  at  cost  to  authors. 
When  proof  is  returned  to  authors,  information 
about  ordering  reprints  will  be  given.  They 
are  obtained  from  Economy  Printing  Co.,  Inc., 
R.D.  3,  Box  169,  Easton,  Maryland  21601-9430. 


MOLLUSK  VOUCHER  SPECIMENS 


It  is  becoming  increasingly  important  for 
future  research  purposes  that  an  identified  sam- 
pling of  species  mentioned  in  publications  be 
deposited  in  a  permanent,  accessible  museum 
specializing  in  mollusks.  This  is  particularly  true 
of  mollusks  used  in  physiological,  medical, 
parasitological,  ecological,  and  experimental 
projects. 


Several  museums  of  natural  history  have  ex- 
tensive modern  facilities  and  equipment  for  the 
housing  and  curating  of  voucher  specimens. 
Material  should  be  accompanied  by  the  identifi- 
cation, locality  data  and  its  bibliographic 
reference.  There  is  no  charge  for  this  perma- 
nent curating  service,  and  catalog  numbers,  if 
desired,  will  be  sent  to  authors  prior  to  publica- 
tion. 


WANTED  -  OLD  SHELL  BOOKS 

Will  pay  good  prices  for  libraries,  second-  Phone  (1-305-725-2260)  or  write:  R.  Tucker 

hand  books  and  reprints  on  mollusks,  shells  Abbott,  American  Malacologists,  Inc.,  P.O. 

and    conchology.    Back    numbers    of    The  Box  2255,  Melbourne,  FL  32902.  Free  ap- 

Nautilus,  vols.  40-71  wanted,  $1.50  each.  praisals. 


»*  .,-••.  * 


americanmalacologists,  mc. 

PUBLISHERS  OF  DISTINCTIVE  BOOKS  ON  MOLLUSKS 

THE  NAUTILUS  (Quarterly) 

MONOGRAPHS  OF  MARINE  MOLLUSCA 

STANDARD  CATALOG  OF  SHELLS 

INDEXES  TO  THE  NAUTILUS 

{Geographical,  vols  1-90;  Scientific  Names,  vols  61-90) 

REGISTER  OF  AMERICAN  MALACOLOGISTS 


JULY  30,  1986 


THE 


I  Marin*  Bic!^j'':3i  Izbarnor; 
LIBRARY 

AUG    8  1986 


Woods  Hole,  Mass. 


NAUTILUS 


ISSN  0028-1344 


Vol.  100 


No.  3 


A  quarterly 

devoted  to 

malacology  and 

the  interests  of 

conchologists 


Founded  1889  by  Henry  A.  Pilsbry.  Continued  by  H.  Burrington  Baker. 
Editor-in-Chief:  R.  Tucker  Abbott 


EDITORIAL  COMMITTEE 
CONSULTING  EDITORS 


Dr.  William  K.  Emerson 
Department  of  Living  Invertebrates 
The  American  Museum  of  Natural  History 
New  York,  NY  10024 

Mr.  Samuel  L.  B.  Fuller 
1053  Mapleton  Avenue 
Suffield,  CT  06078 

Dr.  M.  G.  Harasewych 
363  Crescendo  Way 
Silver  Spring,  MD  20901 

Mr.  Richard  L  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Los  Angeles  County  Museum  of  Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 


Dr.  Arthur  S.  Merrill 
c/o  Department'  of  Mollusks  ' 
Museum  of  Comparative  Zoology 
Cambridge,  MA"  02138     -  ■-'  -   . 

Dr.  Donald  R.  Moore 
Division  of  Marine  Geology 
School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 

Dr.  G.  Alan  Solem 
Department  of  Invertebrates 
Field  Museum  of  Natural  History 
Chicago,  IL  60605 

Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Gilbert  L.  Voss 
Division  of  Biology 

School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 


EDITOR-IN-CHIEF 

Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
Box  2255,  Melbourne,  FL  32902-2255 

Mrs.  Cecelia  W.  Abbott 
Business  and  Subscription  Manager 
P.O.  Box  2255 
Melbourne,  FL  32902-2255 


Second  Class  Postage  paid  at  Melbourne,  Florida 
and  other  post  offices 


The  Nautilus  (USPS  374-980) 
ISSN  0028-1344 

A  quarterly  magazine  devoted  to  malacology. 
Copyright  ^1986  by  American  Malaeolopisls.  Inc. 

OFFICE  OF  PUBLICATION 

American  Malacologists,  Inc.  (United  Parcel  Address: 

2208  South  Colonial  Drive,  Melbourne,  FL  32901) 

Mail:  Box  2255,  Melbourne,  FL  32902-2255 

POSTMASTER:  Send  address  changes  to  above. 


Subscription  Price: 


$15.00  (see  inside  back  cover) 
$17.00  (foreign);  institutions  $20.00 


Marin*  SWofit--*?  Li^WlWY 

LIBRARY 

AUG    8  1986 


Woo<^s  Hole.  Mass. 
CONTENTS^' 


THE 
NAUTILUS 

Volume  100,  number  3  -  July  30,  1986 

ISSN  0028-1344 


M.  G.  Harasewych  and  Richard  E.  Petit 

Notes  on  the  Morphology  of  Admete  viridula  (Gastropoda:  Cancellariidae) . 


.85 


Ian  Stupakoff 

Observations  on  the  Feeding  Behavior  of  the  Gastropod  Pleuroploca  princeps 
(Fasciolariidae)  in  the  Galapagos  Islands 


.92 


William  K.  Emerson 

A  New  Species  of  Morum  from  the  Andaman  Sea  (Gastropoda:  Volutacea). 


.96 


Antonio  J.  Ferreira 

A  New  Deep- Water  Species  of  Lepidoplewrus  (Polyplacophora)  from  the  Venezuela  Basin 98 

William  K.  Emerson  and  Walter  E.  Sage  III 

A  New  Species  of  Lyria  (Gastropoda:  Volutidae)  from  the  Arabian  Sea 101 


David  Nicol  and  Douglas  S.  Jones 

Litharca  lithodomus  and  Adaptive  Radiation  in  Arcacean  Pelecypods. 


.105 


W.  Labato  Paraense 

The  Radula  of  Acrorhis  petricola  (Pulmonata:  Planorbidae) . 


,109 


Harald  A.  Rehder 

Natica  (Glyphepthema)  tedbayeri,  a  replacement  name . 


112 


Book  Reviews 91,  104,  113,  114 


NEW  MANUSCRIPTS 

Contributors  to  The  NaMtilus  are  requested  to  send  their  manuscripts  in  the 
future  to  Dr.  M.  G.  Harasewych,  Editor,  The  Nautilus.  P.  0.  Box  3430,  Silver 
Spring,  MD  20901-0430.  The  list  of  Consulting  Editors  will  remain  largely  the 
same,  and  Dr.  Abbott  will  continue  as  an  Associate  Editor  beginning  with 
volume  101. 


A  HISTORY  OF 
SHELL  COLLECTING 


S.  Peter  Dance 


In  addition  to  being  an  historical 
classic,  this  book  answers  a 
thousand  trivia!  questions: 


A  NEW  AND  ESSENTIAL 
SOURCE  BOOK 

An  intriguing  history  of  shell  collecting  and 
conchology,  from  pre-historic  times  up  to  the 
twentieth  century.  A  scholarly  and  exciting 
description  of  early  expeditions,  infamous  shell 
collectors  of  the  past,  and  the  story  of  the  strug- 
gles and  triumphs  of  early  conchologists.  This 
newly  illustrated  and  expanded  book,  with  a 
more  complete  bibliography,  with  an  enlarged 
American  section,  and  with  an  appendix  to  the 
whereabouts  of  over  600  famous,  old  shell  collec- 
tions, is  a  continuation  of  Peter  Dances'  well- 
known  history  of  shell  collecting. 

I'uhlishi-cl  in  IDKK  l.y  K   J    Hrill  ,.f  Hull.iii.l  ami  iliMnlmU'il  in  North  Ami-rita  iiy 

AMERICAN  MALACOLOGISTS 

(New  Englanii  Division) 

P.O.  Box  1192 

Burlington,  Mass.  0180;Miy2 

280  pp.,  32  pis.  (1  in  color),  text  figs.  Hardbound. 
U.S.  $39.50. 


Who  invented  the  words  concholofjy  and  malacology,  and 
when? 

How  did  Capt.  James  Cooke's  ship  collect  the  first  known 
Sunburst  .Shell  in  New  Zealand? 

What  French  conchologist  pawned  the  family  silverware 
in  order  to  buy  a  rare  Spondylus  Thorny  Oyster? 

What  happened  to  the  valuable  shell  collection  of  the  widow 
of  Capt.  Bligh  of  mutiny  fame? 

How   was  Linnaeus  duped   into  naming  a  common  acid- 
treated  cowrie  as  a  new  species? 

What   beautiful  English  Duchess  with  the  greatest  shell 
collection  used  to  entertain  King  George  111  at  breakfast? 

And  what  ancient  Roman  author  gave  us  the  words,  Tellina, 
I'urjmra  and  Nerila? 

Who  has  the  world's  record  for  collecting  the  most  new 
.species  of  marine  and  land  shells? 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      85 


NOTES  ON  THE  MORPHOLOGY  OF  ADMETE  VIRIDULA 
(GASTROPODA:  CANCELLARHDAE) 


M.  G.  Harasewych  and  Richard  E.  Petit' 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 

Smithsonian  Institution 

Washington,  DC  20560 

ABSTRACT 
The  anatomy  and  shell  morphology  o/ Admete  viridula  (Fabricius,  1780),  a 
boreal  cancellariid,  are  described.  This  species,  the  type  o/ Admete  Kroyer,  18^2, 
differs  from  members  of  the  Cancellariinae  and  Trigonostominae  in  having  a 
thinner  shell  composed  of  a  single  aragonitic  layer,  a  reduced  kidney  and  jaw,  and 
a  long,  expanded  prostate  gland,  as  well  as  in  lacking  a  radula,  separate  sperm 
ingesting  gland  and  dorsally  recurved  albumen  gland.  Based  on  cladistic  analysis 
of  20  shell  and  anatomical  characters,  we  suggest  that  the  family  Admetinae  is  an 
early  offshoot  from  primitive  cancellariid  stock,  that  the  subfamily  Trigono- 
stominae contains  the  most  primitive  cancellariids  studied  anatomically  to  date, 
and  that  Cancellariinae  comprise  a  comparatively  recent  radiation  with 
specialized  chemosensory  capabilities. 


The  composition  and  taxonomic  history  of  the 
superfamily  Cancellariacea  have  been  briefly 
discussed  in  previous  papers  (Harasewych  & 
Petit,  1982,  1984).  The  relationship  of  this  group 
to  other  prosobranchs  is  still  uncertain,  as  is  the 
phylogenetic  arrangement  of  the  nearly  100 
nominal  supraspecific  taxa  proposed  almost  ex- 
clusively on  the  basis  of  shell  characters. 

Recent  supraspecific  taxa  have,  in  general, 
been  clustered  around  three  morphological 
types,  represented  by  the  genera  Cancellaria. 
Trigonostoma  and  Admete,  that  have  been  given 
subfamily  status  by  Cossmann  (1899).  Members 
of  the  Cancellariinae  and  Trigonostominae  in- 
habit tropical  and  temperate  waters  while 
Admetinae  is  generally  limited  to  polar  regions 
with  some  species  inhabiting  deeper  waters  of 
the  temperate  zone.  The  gross  anatomy  of 
Cancellaria  reticulata  (Linne,  1767),  the  type  of 
Cancellariinae,  and  Olssonella  smithii  (Dall, 
1888)  a  trigonostomine,  have  previously  been 
described  (Harasewych  and  Petit,  1982,  1984). 

No  general  anatomical  studies  of  any  adme- 
tine  are  known  to  us,  although  brief  descriptions 
of  the  animal  of  Admete  viridula  (Fabricius, 
1780)  were  given  by  Moller  (1842:  88,  as  A. 


'Research  Associate.  P.O.  Box  30,  North  Myrtle  Beach,  SC 
29582. 


crispa  Moller)  and  Jeffreys  (1876:  322).  Simple 
drawings  of  the  living  animal  were  published  by 
H.  &  A.  Adams  (1853:  pi.  29,  fig.  5;  copied  by 
Tryon,  1885:  pi.  7,  fig.  32),  Troschel  (1865:  pi.  4, 
fig.  14)  and  Morse  (1921:  pi.  7,  fig.  43).  Troschel 
(1865:  pi.  4,  figs.  16,  17)  figured  the  jaws  of  A. 
viridula  and  the  synonymous  A.  crispa,  refer- 
ring to  these  structures  as  "hard  apparatus"  and 
suggesting  that  they  may  be  toxoglossan 
radular  teeth.  The  latter  figure  was  copied  by 
Fischer  (1883:  595,  fig.  359)  and  labelled  as  a 
radular  tooth.  Tryon  (1885:  pi.  7,  fig.  33)  copied 
Fischer's  figure  without  comment.  Thiele  (1904: 
172)  identified  Troschel's  figures  as  jaws  and 
reported  the  absence  of  a  radula  in  A.  viridula. 
The  austral  species  A.  magellanica  Strebel, 
1905,  and  Waipaoa  marwicki  Dell,  1956,  were 
shown  to  lack  radulae,  respectively,  by  Powell 
(1951:  167)  and  Beu  and  Climo  (1974:  327).  Not 
all  admetines  are  without  radulae  as  evidenced 
by  the  figures  of  the  radula  of  Nothoadmete 
tumida  Oliver,  1982,  that  accompanied  the 
description  of  that  species.  The  gross  anatomy 
and  shell  morphology  of  Admete  viridula 
(Fabricius,  1780),  the  type  species  of  Admete 
Moller,  1842  [as  A.  crispa  Moller,  1842]  are  here 
described  and  compared  to  other  cancellariid 
taxa. 
The  checkered  nomenclatural  history  of  this 


86      THE  NAUTILUS 


July  30,  1986 


Vol.  100  (3) 


species  is  detailed  by  Bouchet  &  Waren  (1985: 
257),  who  point  out  that  North  American 
authors  have  used  the  junior  synonym  Admete 
couthouyi  (Jay,  1839)  for  the  species,  based  on 
incorrect  statements  by  Dall  (1887:  298;  1918: 
318,  328)  contending  that  the  type  of  ^4.  viridula 
was  a  turrid.  Although  this  was  refuted  by 
Pilsbry  (1938),  Macpherson  (1971:  107),  citing 
Dall,  followed  other  New  World  malacologists  in 
using  j4.  couthouyi. 

Admete  viridula  is  exceedingly  variable  as  at- 
tested by  the  synonymy  listed  by  Bouchet  & 
Waren  (1985:  258),  to  which  additional  nomina 
will  eventually  be  added.  While  the  Canadian 
specimens  used  for  this  study  are  not  "typical"  of 
the  species,  in  conchological  characters,  they  do 
fall  within  the  range  of  variation  accepted  by 
Bouchet  &  Waren  (1985:  figs.  683-689)  who  ex- 
amined thousands  of  North  Atlantic  specimens. 

Materials  and  Methods 

Anatomical  data  is  based  on  20"  and  19 
specimens  from  the  Saguenay  River,  Quebec 
[48°24'48"N,  70°44'-70°48'W]  Voucher  Material- 
National  Museum  of  Natural  Sciences,  Ottawa 
43946. 

Shell  fragments  for  ultrastructural  studies 
were  broken  from  the  outer  lip  with  pliers. 
Specimens  were  then  immersed  in  10%  hydro- 
chloric acid  (HCl)  until  the  shells  dissolved.  Soft 
parts  were  rinsed  in  distilled  water  and  returned 
to  70%  ethanol  for  dissection.  Portions  of  the 
reproductive  systems  and  anterior  proboscis 
were  sectioned  at  a  thickness  of  6  ^m,  and  the 
sections  stained  in  hematoxylin  and  eosin.  Dried 
shell  fragments  were  powdered  in  a  mortar  and 
pestle,  and  the  X-ray  diffraction  pattern  deter- 
mined on  a  Philips  APD  3600  Automated  X-ray 
Powder  Diffractimeter. 

Shell  Morphology 

External:  Shell,  reaching  20  mm  in  length, 
thin,  ovate  with  conical  spire  and  rounded 
anterior  (Fig.  1).  Protoconch  (Figs.  3,  4)  pauci- 
spiral,  consisting  of  1  whorl,  with  a  smooth, 
chalky  surface  incised  by  fine,  spiral  lines.  Tran- 
sition to  teleoconch  marked  by  beginning  of 
axial  growth  striae  and  an  increase  in  shell 
thickness.  Teleoconch  with  up  to  5V2  convex 
whorls.  Early  whorls  (Fig.  2)  strongly  shoul- 
dered, becoming  more  rounded  with  increase  in 


shell  size.  Suture  deeply  impressed.  Spiral 
sculpture  of  16-21  shallow,  rounded  cords  on 
body  whorl  and  7-10  on  the  penultimate  whorl. 
Axial  sculpture  of  14-24  ribs  per  whorl,  increas- 
ing in  number  but  decreasing  in  prominence  as 
the  shell  becomes  larger.  Aperture  elliptical, 
deflected  from  coiling  axis  by  20-28°.  Outer  lip 
smooth  to  slightly  corrugated.  Siphonal  canal 
short  but  pronounced  in  juveniles  (Fig.  2),  com- 
monly indiscernible  in  large  adults  (Fig.  1). 
Inner  lip  with  2  weak  columellar  folds  and  a 
siphonal  fold.  Color  white  to  ivory,  uniform 
within  and  without. 

Internal:  Fractured  shells  revealed  the  inter- 
nal surfaces  to  be  smooth  and  continuous,  and  to 
lack  the  apertural  lirae  and  periodic  increases  in 
prominence  of  columellar  folds  found  in  other 
cancellariids  (Harasewych  and  Petit,  1982, 
1984). 

Ultrastructure:  A  thin,  brownish,  lamellose 
periostracum  overlies  the  shell,  which  appears 
to  consist  of  a  single  layer  of  crossed-lamellar 
crystals  (Fig.  5),  oriented  with  lamellar  planes 
parallel  to  the  outer  lip.  X-ray  diffraction 
analysis  of  powdered  shell  revealed  it  to  consist 
primarily  (>95%)  of  aragonite. 

Soft-parts  Morphology 

External  features:  The  soft-parts  comprise  3 
to  3V3  whorls,  of  which  the  digestive  gland  (Fig. 
7,  dg)  occupies  about  2V4  whorls,  the  kidney 
(Fig.  7,  k)  less  than  Vs  whorl  and  the  mantle  cavi- 
ty just  over  V2  whorl.  Preserved  animals  were 
orange  tan  in  color,  lacked  any  discernible 
markings  and  were  retracted  at  most  'A  whorl 
into  the  aperture.  The  foot  is  broad  and  ovate, 
lacking  an  operculum.  Tentacles  (Fig.  7,  t)  are 
long,  tubular  and  symmetrical.  The  mantle  edge 
is  smooth,  the  siphon  (Fig.  7,  s)  short  but 
distinct. 

Mantle  rarity:  The  mantle  cavity  is  shorter 
and  broader  than  in  other  cancellariids  dis- 
sected, with  pallial  organs  situated  as  in  other 
higher  prosobranchs.  The  osphradium  (Fig.  7, 
os)  is  broad  anteriorly  (L/W  =  3),  tapers 
posteriorly,  and  consists  of  about  35  leaflets  per 
side.  Adjacent  is  the  ctendium  (Fig.  7,  ct),  twice 
as  long  and  slightly  narrower  than  the 
osphradium,  composed  of  about  70  triangular 
leaflets.  This  organ  and  the  voluminous,  trans- 
versely pleated  hypobranchial  gland  (Fig.  7,  hg) 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      87 


FIGS.  1-6.  Features  of  the  shells  and  jaw  of  Adniete  viridula  (Fabricius).  1,  Apertural  and  right  side  views  of  specimen 
collected  in  the  Saguenay  River,  Quebec.  [48°24'48"N,  70°44'-70°48'W]  sta.  62  30  (NMNS  43946)  3.0  x  .  2,  Scanning  elec- 
tron micrograph  of  juvenile  specimen  dredged  in  673  m,  off  Martha's  Vmeyard,  Massachussetts,  U.S.  Fish.  Comm.  sta. 
994-97.  (USNM  43232)  20.0  x  .  3,  Protoconch  of  specimen  in  figure  2,  lateral  view,  scale  bar  =  200  ^jm.  4,  Protoconch  of 
specimen  in  figure  2,  axial  view,  scale  bar  =  200  /jm.  5,  Fracture  surface.  Plane  of  fracture  parallel  to  outer  lip,  scale  bar 
=  100  fjm.  6,  Lateral  view  of  jaw.  scale  bar  =  50  jjm. 


88      THE  NAUTILUS 


Vol.  100(3) 


vd  pap 


rmc 


FIGS.  7-10.  Anatomical  features  of  Admete  oiridula.  7,  Female  specimen  removed  from  shell,  partially  un- 
coiled and  mantle  cavity  opened  mid-dorsally  to  display  contents.  8,  Dissection  of  anterior  portion  of  proboscis, 
opened  mid-dorsally.  9,  Diagrammatic  representation  of  the  alimentary  system.  10,  Male  reproductive  system. 

a,  anus;  ag,  albumen  gland;  asg,  accessory  salivary  gland;  be,  bursa  copulatrix;  bg,  buccal  ganglia;  bm,  buccal 
mass;  eg,  capsule  gland;  ct,  ctenidium;  dg,  digestive  gland;  hg,  hypobranchial  gland;  int,  intestine;  j,  jaw;  k, 
kidney;  m,  mouth;  me,  mid-esophagus;  ng,  nephridial  gland;  nr,  nerve  ring;  opm,  opening  to  mantle  cavity;  os, 
osphradium;  ot,  oral  tube;  pap,  papilla;  pc,  pericardium;  pen,  penis;  pr,  prostate  gland;  r,  rectum;  rmc,  rear  of 
mantle  cavity;  s,  siphon;  sg,  salivary  gland;  sto,  stomach;  sv,  seminal  vesicle;  t,  tentacle;  tes,  testes;  vd,  vas 
deferens;  vl,  valve  of  Leiblein. 


span  the  roof  of  the  mantle  cavity.  The  pallial 
gonoducts  and  rectum  (Fig.  7,  r)  are  located 
along  the  right  side  of  the  mantle  cavity,  while 
its  rear  is  formed  by  the  pericardium  (Fig.  7,  pc) 
and  kidney  (Fig.  7,  k).  The  kidney  is  small,  being 
only  twice  the  size  of  the  pericardium.  The 
nephridial  gland  (Fig.  7,  ng)  occupies  nearly  half 
the  kidney  along  the  pericardium. 

Alimentary  system:  The  broad,  tubular  pleu- 
rombolic  proboscis  can  be  extended  nearly  the 
length  of  the  shell  aperture,  and  is  the  same 
color  as  the  foot.  Broad  retractor  muscles  ex- 
tend from  the  columellar  muscle,  while  thinner 
strands  originate  from  the  body  wall.  A  short 
oral  tube  (Fig.  8,  ot)  extends  from  the  mouth 
(Fig.  8,  m)  to  the  cuticularized,  tubular  portion 
of  the  jaw  (Fig.  8,  j),  which  expands  posteriorly 
to  envelope  the  lateral  surfaces  of  the  minute 
buccal  mass  (Fig.  8,  bm).  The  longitudinally 


grooved  buccal  mass  lacks  a  radula  and  has  only 
a  vestigial  subradular  membrane.  A  pair  of 
short,  extremely  thin  accessory  salivary  glands 
(Fig.  8,  asg)  lie  alongside  the  buccal  mass, 
emptying  into  the  posterior  portion  of  the  oral 
tube  through  fine  ducts.  The  wider  and  much 
longer  salivary  glands  (Fig.  8,  sg)  join  the  buccal 
mass  laterally,  via  short  ducts,  just  anterior  to 
the  esophageal  opening.  A  small  valve  of 
Leiblein  (Fig.  8,  vl)  and  a  pair  of  buccal  ganglia 
(Fig.  8,  bg)  are  situated  at  the  rear  of  the  bucal 
mass.  The  narrow  mid-esophagus  (Figs.  8,  9, 
me)  runs  posteriorly  from  the  valve  of  Leiblein, 
passing  through  the  nerve  ring  (Fig.  9,  nr) 
before  expanding  and  becoming  sacular.  The 
esophagus  narrows  at  the  rear  of  the  cephalic 
sinus  and  leads  to  a  simple  U-shaped  stomach 
(Fig.  7,  9,  sto)  that  is  embedded  in  the  anterior 
face  of  the  digestive  gland.  Longitudinal  folds 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      89 


TABLE  1.  Admete  rindutn  (Fabricius),  measurements  of 
shell  characters.  Linear  measurements  in  millimeters,  (n  = 
10). 

Standard 
Character  Mean        Deviation  Range 


shell  length 

16.87 

1.91 

13.82-19.02 

shell  width 

9.90 

1.13 

8.18-10.97 

aperture  length 

10.21 

1.42 

8.13-11.58 

aperture  length 

0.604 

0.023 

0.567-0.641 

shell  length 

#whorls,  teleoconch 

4.78 

0.35 

4.33-5.33 

spire  angle 

.5.5.2° 

2.9° 

45.5°-.58.0° 

line  the  stomach  and  pass  into  the  intestine  (Fig. 
9,  int),  which  runs  through  the  kidney  and  along 
the  right  wall  of  the  mantle  cavity,  expanding 
slightly  to  form  the  rectum  (Figs.  7,  9,  r). 
Neither  an  anal  gland  nor  identifiable  gut  con- 
tents were  detected  in  any  of  the  specimens  ex- 
amined. 

Female  reproductive  system:  As  the  only 
female  specimen  available  to  us  was  poorly  pre- 
served, only  the  pallial  portion  of  the  oviduct  is 
described.  The  oviduct  passes  through  the 
kidney  and  expands  to  form  a  broad,  laterally 
compressed  albumen  gland  (Fig.  7,  ag),  which 
expands  further  to  form  the  long,  blunt,  capsule 
gland  (Fig.  7,  eg).  A  small,  ovate,  bursa  copula- 
trix  (Fig.  7,  be)  is  situated  below  the  anterior 
margin  of  the  capsule  gland.  An  ingesting  gland 
was  not  found.  Egg  capsules  oi  Admete  viridula 
are  figured  by  Bouchet  and  Waren  (1985:  fig. 
687). 

Male  reproductive  system:  The  testes  (Fig.  10, 
te)  consist  of  a  yellowish  mass  of  ascinous 
tubules  embedded  in  the  right  side  of  the  diges- 
tive gland  and  extend  nearly  a  full  whorl  from 
its  anterior  edge.  These  tubules  converge  to 
form  a  testicular  duct  that  immediately  forms  a 
serpentine  seminal  vesicle  (Fig.  10,  sv).  The  duct 
straightens  at  the  anterior  limit  of  the  digestive 
gland,  passes  through  the  kidney  and  enters  the 
mantle  cavity  ventral  to  the  rectum.  It  greatly 
expands  to  form  the  prostate  gland  (Fig.  10,  pr), 
which  extends  along  the  posterior  third  of  the 
mantle  cavity  and  communicates  with  it  by  a 
long  ventral  slit  (Fig.  10,  opm).  From  the 
anterior  portion  of  the  prostate  gland,  a  tubular 
vas  deferens  (Fig.  10,  vd)  leads  to  the  base  of  the 
penis  (Fig.  10,  pen),  which  is  ovate  in  cross- 
section,  extends  approximately  %  the  length  of 
the  mantle  cavity  and  has  a  terminal  papilla 
(Fig.  10,  pap). 

Nervous  system:  Admete  viridula  has  a  typical 


cancellariid  nervous  system  (Bouvier,  1887: 
Harasewych  and  Petit,  1982,  1984),  with  the 
cerebral,  pedal  and  pleural  and  subesophageal 
ganglia  fused  into  a  nerve  ring,  and  the  buccal 
ganglia  (Fig.  8,  bg)  at  the  rear  of  the  buccal 
mass. 

Discussion 

Cladistic  analysis  of  the  20  shell  and 
anatomical  characters  listed  in  Table  2  produced 
the  cladogram  in  Figure  11.  Despite  the  lack  of 
agreement  on  the  relationships  of  supraspecific 
taxa,  nearly  all  classifications  proposed  to  date 
separate  the  Admetinae  from  the  remaining 
Cancellariidae  (H.  and  A.  Adams,  1853: 
Troschel,  1865;  Fischer,  1883;  Tryon,  1885; 
Cossmann,  1899;  Thiele,  1929;  Wenz,  1943),  and 
this  arrangement  is  supported  by  the  present 
study. 

Of  the  characters  used  to  distinguish  the 
Admetinae  from  other  cancellariids,  most  con- 

TABLE  2.  Shell  and  anatomical  characters  used  to  distin- 
guish between  Admete  viridula,  Cancellaria  reticulata  and 
Olssonella  smithii.  Primitive  states  denoted  by  (0),  derived 
states  by  (1). 

1)  Shell:  (0)  thick:  (1)  thin. 

2)  Internal  varices:  (0)  pronounced;  (1)  reduced  or  absent. 

3)  Shell  ultrastructure  of:  (0)  2  orthogonal  layers;  (1)  1 
layer. 

4)  Protoconch  of:  (0)  2  whorls;  (1)  1  whorl. 

5)  Tentacles:  (0)  symmetrical,  unmodified;  (1)  asymmetri- 
cal, left  tentacle  modified. 

6)  Osphradium  with  approximately:  (0)  35  leaflets/side;  (1) 
70  leaflets/side. 

7)  Ctenidium  with:  (0)  <  125  leaflets;  (1)  >  200  leaflets. 

8)  Distance  between  osphradium  and  ctenidium:  (0)  nor- 
mal; (1)  large  relative  to  either  organ. 

9)  Kidney  occupying;  (0)  >  'A  whorl;  (1)  <  Vs  whorl. 

10)  Proboscis:  (0)  tubular;  (1)  ventrally  flattened,  papillose. 

11)  Jaws  with:  (0)  short  posterior  lobes;  (1)  long  posterior 
lobes. 

12)  Buccal  mass:  (0)  small;  (1)  large,  filling  retracted  pro- 
boscis. 

13)  Radula:  (0)  present;  (1)  absent. 

14)  Outer  cusps  of  radular  teeth:  (0)  simple,  smooth;  (1) 
with  secondary  dentition. 

15)  Accessory  salivary  gland:  (0)  shorter  than;  (1)  longer 
than;  salivary  gland. 

16)  Mid-esophagus  posterior  to  nerve  ring:  (0|  sacular;  (1) 
convoluted  tube. 

17)  Anal  gland:  (0)  present;  (1)  absent. 

18)  Sperm  ingesting  gland:  (0)  present;  (1)  absent. 

19)  Albumen  gland:  (0)  dorsally  recurved;  (1)  laterally  com- 
pressed. 

20)  Prostate  gland:   (0)   long,   expanded;   (1)  short,   con- 
voluted. 


90      THE  NAUTILUS 


July  30,  1986 


Vol.  100  (3) 


3 

•o 


> 


CO 

w 

3 

o 
o 
CJ 


E 


FIG.  11.  Cladogram  of  phylogenetic  relationships  of  cancel- 
lariid  taxa.  Single  slashes  across  tree  branches  represent 
transformations  of  the  corresponding  character  from  the 
primitive  (0)  to  the  derived  (1)  state.  Double  slashes  indicate 
transformations  that  occur  more  than  once. 

spicuous  are  their  thin  shells  lacking  pro- 
nounced surface  sculpture  and  apertural  lirae. 
This  is  likely  a  consequence  of  their  polar 
habitat,  as  Graus  (1974)  has  shown  that  de- 
creased calcification  in  high  latitudes  is  due  to 
reduced  availability  of  calcium  carbonate  in 
colder  waters.  Admetines  also  lack  such  anti- 
predatory  features  as  internal  varices  that  are 
found  in  tropical  and  temperate  cancellariids. 
The  predominance  of  aragonite  in  the  shell  is 
plesiomorphic,  as  calcite  is  more  easily  formed 
(Wilbur,  1964)  and  more  stable  (Lowenstam, 
1954)  at  low  temperatures. 

In  terms  of  anatomical  organization,  Admete 
viridula  differs  from  ancestral  cancellariid  mor- 
phology, most  nearly  approximated  by  Olsso- 
nella  snulhii  of  the  taxa  studied  to  date,  in  hav- 
ing a  shallower  mantle  cavity,  a  reduced  kidney, 
and  in  lacking  color  pattern.  At  least  some  of 
these  may  also  be  adaptations  to  cold  water. 

Cancellariines  and  Trigonostomines  have  uni- 
serial  "tricusped"  radular  teeth  with  elaborate 
secondary  dentition  on  the  outer  cusps  and  a 
comparatively  simple  ventrally  recurved  central 
cusp  (Harasewych  and  Petit,  1982,  1984;  Petit 
and  Harasewych,  1986).  The  radula  of  Notho- 


admete  tumida,  the  only  known  radulate 
admetine,  has  barbed  central  cusps  and  simple 
bulbous  outer  cusps  (Oliver,  1982:  figs.  3,  5). 
Presence  of  central  cusp  barbs  in  Cancellaria 
atopodonta  Petit  and  Harasewych  (1986:  figs. 
15,  16)  suggests  that  this  feature  is  primitive 
and  occurred  in  the  ancestor  of  all  Recent 
cancellariids.  The  jaws  of  admetines  lack  the 
long  posterior  lobes  found  in  cancellariines  and 
trigonostomines.  Although  the  diet  of  cancel- 
lariids remains  unknown,  we  have  suggested 
that  they  are  piercing  suctorial  feeders  based  on 
the  functional  morphology  of  their  alimentary 
systems  (Harasewych  and  Petit,  1982,  1984; 
Petit  and  Harasewych,  1986).  The  loss  of  radula 
and  reduction  in  the  size  of  the  buccal  mass  in 
most  admetines  suggests  that  they  feed  suc- 
torially,  but  on  a  different  group  of  prey 
organisms  than  other  cancellariids. 

The  lack  of  an  anal  gland  in  Admete  viridula 
and  Cancellaria  reticulata  is  likely  a  com- 
paratively recent  convergence,  as  this  organ  has 
been  reported  in  Cancellaria  cancellata 
(Graham,  1966). 

Additional  differences  between  admetines  and 
other  cancellariids  are  found  in  the  reproductive 
systems.  The  prostate  gland  ofAdynete  viridula 
is  long  and  greatly  expanded,  while  in  Olssonella 
smithii  and  Cancellaria  reticulata  it  is  short  and 
convoluted.  Admete  viridula  also  lacks  the 
separate  sperm  ingesting  gland  and  dorsally 
recurved  albumen  gland  found  in  cancellariines 
and  trigonostomines. 

The  phylogenetic  arrangement  in  Figure  1 1  is 
supported  by  the  fossil  record,  as  several 
"admetines"  have  been  reported  from  the  Upper 
Cretaceous,  and  Waipaoa.  known  to  be 
aradulate  in  the  Recent  fauna,  dates  back  to  the 
Oligocene.  Cancellaria,  sensu  stricto,  first  ap- 
pears in  the  Miocene  and  is  characterized  by 
modifications  to  the  left  cephalic  tentacle,  pro- 
boscis, osphradium  and  ctendium,  organs  in- 
volved in  tactile  and  distance  chemoreception. 

Acknowledgments 

We  thank  Jane  B.  Topping,  Invertebrate 
Zoology  Division,  National  Museum  of  Natural 
Sciences,  Ottawa,  for  making  available  the 
preserved  specimens  used  in  this  study.  Some  of 
this  work  was  done  at  the  Smithsonian  Marine 
Station  at  Link  Port,  Florida,  and  we  thank  Dr. 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      91 


Mary  Rice  for  making  these  facilities  available. 
This  is  contribution  number  158  of  the  Smith- 
sonian Marine  Station  at  Link  Port.  Critical 
review  of  the  manuscript  by  Dr.  R.  S.  Houbrick, 
National  Museum  of  Natural  History,  Smith- 
sonian Institution,  is  gratefully  acknowledged. 

LITERATURE  CITED 

Adams,  H.  and  A.  Adams.  1853-58.  The  Getiera  of  Recent 

MoUusca.  3  vols.  London. 
Beu,  A.  G.  and  F.  M.  Climo.    1974.  Mollusca  from  a  recent 

coral  community  in  Palliser  Bay,  Cook  Strait.  N.  Z.  Joum. 

Marine  and  Freshwater  Research  8(2):307-332. 
Bouchet,  P.  and  A.  Waren.  1985.  Revision  of  the  Northeast 

Atlantic  Bathyal  and  Abyssal  Neogastropoda  excluding 

Turridae    (Mollusca,    Gastropoda).    Boll.    Malacologico. 

Suppl.  1:121-296. 
Bouvier,    E.    L.    1887.    Systeme    nerveux.    morphologie 

generale  et  classification  des  Gasteropodes  prosobranches. 

Ann.  Sci.  nat.  Zool.  3:1-510. 
Cossmann,   M.   1899.  Caneellariidae.  Essais  de  Paleocon- 

chologie  Ccmiparee  3:1-41,  pis.  1-2. 
Dall,  W.  H.  1887.  Supplementary  notes  on  some  species  of 

mollusks  of  the  Bering  Sea  and  vicinity.  Proc.  U.S.  Nat. 

Miis.  9:297-309,  pis.  3-4. 
1888.  [in]  Agassiz,  A.  Three  Cruises  of  the  United 

States  Coast  and  Geodetic  Survey  Steamer  Blake,  2(8): 

62-75,  figs.  282-312. 
1918.  Notes  on  the  nomenclature  of  the  mollusks 

of  the  family  Turridae.  Proc.  U.S.  Nat.  Mus.  54:313-333. 
Dell,  R.  K.  1956.  The  Archibenthal  Mollusca  of  New  Zealand. 

Dominion  Mus.  Bull.  18:1-235,  pis.  1-25,  A,  B. 
Fabricius,  0.  1780.  Fauna  Groenlandica.  Hafnia  et  Lipsiae. 

452  p. 
Fischer,    P.    1880-1887.   Manuel   de  conchyhologie   et   de 

paleontologie  conchyliologique  ou  histoire  naturelle  des 

moUusques  vivants  etfossil.es.  Paris,  F.  Savy.  1369  p. 
Graham,  A.  1966.  The  fore-gut  of  some  marginellid  and  can- 

cellariid    prosobranchs.    Stud.    trop.    Oceanogr.    Miami 

4:134-151. 
Graus,  R.  R.  1974.  Latitudinal  trends  in  the  shell  character- 
istics of  marine  gastropods.  Lethaia  7:303-314. 
Harasewych,  M.  G.  and  R.  E.  Petit.  1982.  Notes  on  the  mor- 
phology of  Cancellaria  reticulata  (Gastropoda:  Caneellari- 
idae). The  Nautilus  96(3):104-113. 
1984.  Notes  on  the  morphology  of  Olssonella 

smithii    (Gastropoda:    Caneellariidae).     The    Nautilus 

98(l):37-44. 


.Jay,  J.  C.  1839.  A  catalogue  of  the  shells  .  .  .  contained  in  the 

collection  of  John  C.  Jay.  M.D.  Ed.  3,  New  York.  125  p., 
10  pis. 
Jeffreys,  J.  G.  1876.  New  and  peculiar  mollusca  of  the  Euli- 

midae  and  other  families  of  gastropoda,  as  well  as  the 

pteropoda,  procured  in  the  'Valerous'  expedition.  Ann. 

Mag.  nat.  Hist.  4(19):317-339. 
Linne,    C.    von.    1767.   Systema    naturae   per   regna    tria 

naturae.     Editio    duodecimo    reformata.     Vol.     1     (2): 

533-1327.  Stockholm. 
Lowenstam,  H.  A.  1954.  Factors  affecting  the  aragonite: 

calcite  ratios  in  carbonate  secreting  marine  organisms.  J. 

Geol.  62:284-322. 
Macpherson,    E.    1971.    The    Marine    Molluscs   of   Arctic 

Canada.  Nat.  Mus.  Canada,  Publ.  in  Biol.  Oceanography, 

No.  3:1-149. 
Moller,  H.  P.  C.   1842.  Index  Molluscorum  Groenlandiae. 

Nat.  Tidsskr.  4:76-97. 
Morse,  E.  S.  1921.  Observations  on  living  gastropods  of  New 

England.  Peabody  Museum,  Salem.  29  pp.,  9  pis. 
Oliver,  P.  G.  1982.  A  new  species  of  Cancellariid  gastropod 

from  Antarctica  with  a  description  of  the  radula.  Br. 

Antarct.  Surv.  Bull.  57:15-20. 
Petit,  R.  E.  and  M.  G.  Harasewych.  1986.  New  Philippine 

Caneellariidae  (Gastropoda:  Cancellariacea),  with  notes  on 

the  fine  structure  and   function  of  the  nematoglossan 

radula.  The  Veliger  28(4):436-443. 
Pilsbry,  H.  A.  1938.  On  the  history  and  status  oi Lora  Gistel. 

The  Nautilus  51(4):11.5-118. 
Powell,  A.  W.  B.  1951.  Antarctic  and  Subantarctic  Mollusca: 

Pelecypoda  and  Gastropoda.  Discovery  Repts.  26:47-196, 

pis.  5-10. 
Strebel,   H.    1905.   Beitrage  zur  Kenntnis  der  Mollusken 

Fauna  der  Magalhaen-Provinz.  Zool.  Jb.,  Abt.  Syst.,  Jena 

22:575-666,  pis.  21-24. 
Thiele,   J.    1903  [1904].   Die  beschalten  Gastropoden  der 

deutschen  Tiefsee-Expedition  1898-1899.  B.  Anatomisch- 

systematische    Untersuchungen    einiger    Gastropoden. 

Deutsche  Tiefsee-Exped..  Bd.  VII:149-179,  Taf.  1-9. 
1929-35.  Handbuch  der  system.ati.'ichen  Weich- 

tierkunde.  Jena,  Gustave  Fischer.  1154  p. 
Troschel,  F.  H.  1856-1893.  Das  Gebiss  der  Schnecken.  zur 

Begriindung  einer  natilrlichen  Classification.  2.  Berlin. 
Tryon,  G.  W.  1885.  Family  Caneellariidae.  Manual  ofCon- 

chology  7:65-98,  pis.  1-7. 
Wenz,  W.  1938-1943.  Handbuch  der  Paldozoologie  (0.  H. 

Schindewolf,  ed.),  Berlin,  Band  6,  Teil  1  (pt.  2)  1639  p. 
Wilbur,  K.  M.  1964.  Shell  formation  and  regeneration  [in] 

Physiology  of  Mollusca,  vol.  1,  Wilbur,  K.  M.  and  C.  M. 

Yonge  eds.  Academic  Press,  New  York. 


BOOK  REVIEW 


Washington  Public  Shore  Guide:  Marine  Waters 
by  James  W.  Scott  and  Melly  A.  Reuling. 
1986.  348  pp.,  numerous  photos,  maps  and 
habitat  sketches.  Cloth,  $25.00;  paperback, 
$14.95. 

If  you  are  planning  on  studying  or  visiting  the 


shorelines  of  the  State  of  Washington,  this  is  a 
great  bargain  with  many  useful  facts  about  the 
2,400-mile  saltwater  shoreline  of  14  counties.  It 
largely  ignores  mollusks,  a  fact  which  offers  a 
challenge  to  conchologists.-7?.  T.  Abbott. 


92      THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


OBSERVATIONS  ON  THE  FEEDING  BEHAVIOR  OF  THE  GASTROPOD 

PLEUROPLOCA  PRINCEPS  (FASCIOLARIIDAE)  IN 

THE  GALAPAGOS  ISLANDS 

Ian  Stupakoff 

Department  of  Invertebrates 

American  Museum  of  Natural  History 

Central  Park  West  at  79th  Street 

New  York,  NY  10024 

ABSTRACT 

The  feeding  behavior  of  the  gastropod.  Pleuroploca  princeps,  (Fasciolariidae)  is 
reported.  Eight  species  of  gastropods  were  found  to  be  preyed  upon  by  P.  princeps; 
two  species  of  crabs  were  eaten  as  carrion.  No  signs  of  cannibalism  were  noticed, 
but  intraspecific  competition  was  observed. 


While  working  at  the  marine  laboratory  of  the 
Charles  Darwin  Research  Station,  Isla  Santa 
Cruz,  Galapagos,  Ecuador,  I  had  the  opportuni- 
ty to  collect  the  gastropod,  Pleuroploca  princeps 
(Sowerby,  1825),  (Keen,  1971,  p.  611,  fig.  1324), 
and  to  study  its  feeding  habits. 

The  only  literature  citation  on  the  feeding 
habits  of  P.  princeps  is  by  Sorensen  (1943,  p.  4), 
who  observed  P.  princeps  feeding  on  Phyllono- 
tus  bicolor  (now  Hexaplex  erythrostomus  (Swain- 
son))  in  Bahia  San  Carlos,  Gulf  of  California. 
Other  observations  on  feeding  habits  are 
reported  for  P.  gigantea  (Kiener)  in  the  western 
Atlantic  by  Menzel  and  Nichy  (1958,  p.  144), 
Paine  (1963  a,  pp.  66,  67;  1963  b,  pp.  402,  403), 
and  Cornellisson  (1985,  p.  125),  and  on  P. 
trapezium  (Linnaeus)  and  P.  filamentosa 
(Roding)  in  the  Indo-Pacific  by  Risbec  (1932,  pp. 
374,  375),  Maes  (1967,  p.  51),  and  Reyes  (1981, 
p.  4).  The  major  prey  items  in  the  diet  of 
members  of  the  family  Fasciolariidae  are  gas- 
tropods, bivalves,  sedentary  polychaetes,  cirri- 
pedes  and  carrion  (Taylor,  Morris  and  Taylor, 
1980,  p.  377,  table  1). 

Pleuroploca  princeps  is  a  characteristic  faunal 
constituent  of  the  Panamanian  province  and  is 
one  of  the  largest  gastropods  in  the  Colon 
Archipelago  (Wellington,  1975,  p.  59),  attaining 
250  mm.  in  length.  This  species  is  widely  distri- 
buted in  the  Galapagos  Islands.  Several 
specimens  were  collected  or  observed  around 
the  islands  of  Santa  Fe,  Santa  Cruz,  Espahola, 
Floreana,  Bartolome,  Cousins  Rock  and  San 
Cristobal.  Living  specimens  were  noted  in  the 


subtidal  zone  and  to  depths  of  21  meters  (per- 
sonal observation).  They  were  found  on  lava 
rocks  or  on  sandy  bottoms  but  occur  more  fre- 
quently in  areas  where  the  lava  boulders  meet 
the  sandy  bottom. 

The  specimens  of  P.  princeps  used  in  this 
study  were  collected  on  lava  boulders  and  sand 
at  Isla  Santa  Fe  (July,  1984)  and  Isla  Santa  Cruz 
(Oct.  and  Nov.  1984).  The  animals  were  kept  in 
aquaria  at  the  marine  laboratory. 

Feeding  Behavior 

Since  P.  princeps  is  a  typical  neogastropod,  it 
feeds  mainly  on  other  gastropods  (Table  1).  Dur- 
ing these  observations  no  bivalves  were  offered 
as  food.  In  aquaria,  P.  princeps  were  observed 
to  feed  on  dead  crabs  such  as  Grapsus  grapsus 
(Linnaeus)  and  Petrolisthes  edwardsii 
(Saussure),  (see  Table  1  and  Fig.  2).  A  specimen 
of  P.  princeps  tried  several  times  to  attack  a 
Nerita  .^cabricosta  (Lamarck)  but  could  not 
reach  the  soft  body  of  the  prey  with  its  proboscis 
because  of  the  protection  of  the  heavily  calcified 
operculum  of  the  nerite. 

When  P.  princeps  was  at  rest,  the  body  and 
half  of  the  shell  was  buried  in  the  sandy  substra- 
tum (personal  observation).  It  was  at  this 
resting  time  that  five  specimens  of  Thais 
melones  (Duclos)  were  introduced  into  the 
aquarium  as  food.  By  using  what  are  thought  to 
be  chemoreceptors  (fide  Kohn,  1961),  the 
predator  detected  the  presence  of  the  prey  and 
began  its  search. 

The  eyes  of  P.  princeps  are  poorly  developed 


Vol.  100(3) 


July  30,  1986  THE  NAUTILUS      93 

TABLE  1.  Species  eaten  by  Plcurophca  princeps  in  aquarium. 


Phylum 

Class 

Faml ly 

Species 

MoIIusca 

Gastropoda 

Cymati  idae 

Cymatium  partheopeum 

Thaididae 

Thais  melones 
Purpura  columellaris 
Purpura  pansa 

Muricidae 

Muricanthus  princeps 

Tonnldae 

Ma  lea  r ingens 

(pers.  obs.  J.  de  Roy) 

Cypraeidae 

Cypraea  cervinetta 

Conidae 

Conus  brunneus 

Arthropoda 

Crustacea 

Grapsidae 

Grapsus  grapsus 

Porcellanidae 

Petrolisthes  edwardsii 

and  probably  cannot  visually  detect  prey.  Conse- 
quently, this  predator  must  feel  its  way  to  the 
prey  with  its  massive  foot  (Fig.  A).  On  contact, 
P.  princeps  wraps  the  prey  in  its  foot  and  begins 
a  series  of  usually  two  or  three  muscular  con- 
tractions as  observed  in  20  cases.  Raising  its 
shell  slowly  and  then  lowering  it  very  quickly 
against  its  prey  (Figs.  B  and  C),  the  predator 
succeeds  in  making  the  prey  withdraw  inside  its 
shell.  When  the  prey  has  withdrawn,  the  shell  is 
maneuvered  until  the  aperture  is  positioned  up- 
ward and  close  to  the  predator's  proboscis.  At 
the  same  time,  the  predator's  muscular  foot 
closes  the  prey's  siphonal  canal  which  may  last 
from  a  few  minutes  up  to  two  days  depending  on 
the  size,  shape  and  strength  of  the  prey  (arrow 
in  Fig.  E).  Figure  1  shows  how  P.  princeps 
envelopes  a  Cymatium.  partheyiopeum  (von 
Salis)  by  the  same  method  described  above. 
When  this  series  of  movements  is  terminated,  P. 
princeps  prys  up  the  prey's  operculum  with  its 
proboscis,  eventually  penetrating  the  prey's 
body  and  devouring  the  soft  parts  (Fig.  E).  The 
whole  animal  is  consumed  and  the  muscular 
tissue  attached  to  the  operculum  is  rasped  clean 
by  radular  action.  Similar  feeding  habits  were 
described  for  Fasciolaria  hunteria  (Perry), 
(Wells,  1958,  p.  155)  and  for  Pleuroploca 
gigantea  (Paine,  1963  a,  p.  67). 

Competition  and  Cannibalism 

In  one  of  the  aquaria,  two  adult  specimens  of 


P.  princeps  were  deprived  of  food  for  a  period  of 
two  weeks.  During  this  time  no  signs  of  attack 
or  cannibalism  were  noticed.  No  references  to 
cannibalism  by  Pleuroploca  have  been  located  in 
the  literature.  In  contrast,  Fasciolaria  tulipa 
(Linnaeus)  (Snyder  &  Snyder,  1971,  p.  257)  and 
F.  hunteria  (Wells,  1958,  p.  155)  will  cannibalize 
even  when  other  sources  of  food  are  available. 
Nevertheless,  after  this  period  of  starvation  one 
specimen  of  T.  melones  was  offered  as  food  to 
these  two  specimens.  At  this  moment,  competi- 
tion was  observed  between  the  two  P.  princeps. 
The  specimen  (approx.  100  mm.)  that  had 
enveloped  the  prey  was  then  attacked  by  the 
other  larger  and  heavier  specimen  (approx.  125 
mm.)  which  "unwrapped"  the  prey  and  removed 
it  from  its  first  predator  in  a  period  of  two 
minutes.  The  smaller  predator  tried  to  recover 
its  prey  four  times  without  success. 

Acknowledgments 

I  thank  the  following  people  for  their 
comments  on  the  manuscript:  Dr.  William  K. 
Emerson  and  Mr.  Walter  E.  Sage,  III,  of  the 
American  Museum  of  Natural  History,  Dr.  Jan 
Pechenick  of  Tufts  University,  and  Dr.  Ruth  D. 
Turner  of  Harvard's  Museum  of  Comparative 
Zoology,  and  to  Dr.  R.  Tucker  Abbott  for  editing 
this  document.  I  am  very  grateful  to  Dr. 
Gunther  Reck,  Director  of  the  Charles  Darwin 
Research  Station,  for  the  wonderful  opportunity 
to  work  on  the  Galapagos  Islands.  Support  in 


94      THE  NAUTILUS 


July  30,  1986 


Vol.  100  (3) 


FIG.  1,  Adult  Pleuroploca  prinreps  (.shell  approx.  12r)  mm.)  ciivflciping  a  live  Cynuitium  parthenopeum  and  closing  the 
prey's  siphonal  canal  with  its  muscular  foot  (see  arrow);  2,  Adult  Pleuroploca  princeps  feeding  on  a  dead  Porcelain  crab. 
Petrolitkes  edwardsii  (natural  size). 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      95 


part  was  received  by  the  Undergraduate- 
Graduate  Research  Program  of  the  American 
Museum  of  Natural  History  supported  by  the 
Greenwall  Foundation. 

LITERATURE  CITED 

Cornellisson,  H.  1985.  Florida  Shelling-Perseverance  pays 
off!  Texas  Conchologist  21(4):124-126. 

Keen,  A.  M.  1971.  Sea  Shells  of  Tropical  West  America. 
(Second  Ed.),  Stanford  University  Press,  Stanford,  CA., 
xiv  +  1064  pp. 

Kohn,  A.  J.  1961.  Chemoreception  in  Gastropod  Molluscs. 
A7yu;rican  Zoologist  l(2):291-308. 

Maes,  V.  0.  1967.  Radulae  of  Two  Species  of  Pleuroploca 
(Fasciolariidae)  From  The  Indo-Pacific.  The  Nautilus 
81(2):48-54. 

Menzel,  R.  W.  &  F.  E.  Nichy.  1958.  Studies  of  The  Distribu- 
tion And  Feeding  Habits  of  Some  Oyster  Predators  in  Alli- 
gator Harbor,  Florida.  Bull.  Mar.  Sci.  Gulf  and  Caribbean 
8(2):125-145. 

Paine,  R.  T.  1963  a.  Trophic  Relationships  of  8  Sympatric 
Predatory  Gastropods.  Ecology  44(l):63-73. 

1963  b.  Feeding  Rate  of  a  Predaceous  Gastro- 
pod, Ple^iroploca  gigantea.  Ecology  44(2):402-403. 

Reyes,  C.  W.  1981.  The  Fascination  of  Live  Shells.  Carfel 
(Manila)  3(5):4,  10. 

Risbec,  J.  1932.  Note  Sur  Les  Moeurs  de  Ricinula  chaidea 
Duel,  et  de  Fasciolaria  filamentosa  Lmk.  Bull.  Sac.  Zool. 
Ft.  57:374-375. 

Snyder,  N.  F.  R,  &  H.  A.  Snyder.  (1971).  Pheromone-Medi- 
ated  Behaviour  of  Fasfio/aj-ia  tulipa.  Anim.  Behav.  19(2): 
257-268. 

Sorensen,  A.  (1943).  Traveling  and  Collecting  in  Mexico.  The 
Nautilus  57(l):l-5. 

Taylor,  J.  D.,  Morris,  N.  .J,  and  C.  N.  Taylor.  1980.  Food 
Specialization  and  the  Evolution  of  Predatory  Proso- 
branch  Gastropods.  Palaeontology  23(2):375-409. 

Wellington,  G.  M.  1975.  Medios  Ambientes  Marinos  Y 
Costeros  de  Galapagos.  Ms.  on  file  in  library  of  the  Charles 
Darwin  Research  Station.  Isla  Santa  Cruz,  Galapagos, 
Ecuador. 

Wells,  H.  W.  1958.  Predation  of  Pelecypods  and  Gastropods 
by  Fasciolarta  huntena  (Perry).  Bull.  Mar.  Sci.  Gulf  and 
Caribbean  8(2):  152- 166. 


FIG.    3    A-E,     Sequence    of    capturing    movements    of 
Pleuroploca  princeps. 


96      THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


A  NEW  SPECIES  OF  MORUM  FROM  THE  ANDAMAN  SEA 
(GASTROPODA:  VOLUTACEA) 


William  K.  Emerson 

Department  of  Invertebrates 

American  Museum  of  Natural  History 

New  York,  NY  10024 

ABSTRACT 
Morum  (Oniscidia)  ninomiyai,  new  species,  is  described  from  off  Thailand  in  the 
Andaman  Sea  and  is  compared  with  closely  related  congeners. 


In  1981,  the  late  Carl  C.  Withrow  of  Florida 
submitted  to  me  for  identification  a  specimen  of 
an  apparently  new  species  of  Morum  from  the 
Andaman  Sea,  off  southern  Thailand.  This 
unique  specimen  was  subsequently  illustrated  by 
Sally  Diana  Kaicher  (1983)  in  her  "Card 
Catalogue  of  World-Wide  Shells",  as  an  ap- 
parently undescribed  Morum.  The  Andaman 
Sea  specimen  was  again  transmitted  to  me  for 
study  in  1983  by  Taizo  Ninomiya  of  Tokyo,  who 
had  obtained  Mr.  Withrow's  collection  of 
Morum.  Mr.  Ninomiya  kindly  permitted  me  to 
retain  the  specimen  with  the  expectation  that 
additional  material  might  be  forthcoming  from 
his  contacts  in  Bangkok.  Fortunately,  a  second 
specimen  resulted  from  the  inquiries  of  Mr. 
Ninomiya,  for  whom  I  take  great  pleasure  in 
naming  this  interesting  discovery. 

The  new  species  is  a  member  of  an  Indo- 
Pacific  group  within  the  subgenus  Oniscidia 
Morch,  1852,  composed  of  the  following  taxa: 
cancellatum  (Sowerby,  1824;  type  species); 
grande  (A.  Adams,  1855);  uchiyamai  Kuroda 
and  Habe,  in  Habe,  \^&\\  joelgreenei  Emerson, 
1981;  and  watanabei  Kosuge,  1981. 

The  genus  Morum  (sensu  lato)  has  long  been 
classified  with  the  mesogastropods  in  the 
Cassidae  (Thiele,  1929;  Boss,  1982;  Emerson, 
1985).  Anatomical  studies  of  Morum,  however, 
indicate  that  this  genus  is  referable  to  the 
neogastropod  family  Harpidae  (Hughes,  1986, 
and  personal  communications).  Dr.  Hughes  has 
in  preparation  a  manuscript  in  which  he  rectifies 
the  present  taxonomic  misplacement  of  the 
genus.  Recent  observations  on  New  and  Old 
World  species  of  Morum  (sensu  stricto  and 
Oniscidia),  moreover,  confirm  that  these 
gastropods  also  autotomize  the  posterior  por- 


tion of  the  foot  (R.  Goldberg,  P.  Williams  and  W. 
Liltved,  personal  communications),  a  well- 
known  behavorial  characteristic  of  the'  genus 
Harpa  (Rehder,  1973). 

It  should  be  noted  that  Harpidae  Bronn,  1849 
(type  genus  Harpa  (Roding,  1798); 
(Gastropoda),  is  a  homonym  of  Harpidae  Hawle 
and  Corda,  1847  (type  genus  Harpes  Goldfuss, 
1839)  (Trilobita).  Raven  (1985)  has  applied  to  the 
International  Commission  On  Zoological 
Nomenclature  to  emend  the  spelling  of  this  gas- 
tropod family  to  Harpaidae  Bronn,  1849  and  to 
place  the  emended  taxon  on  the  Official  List  of 
Family-Group  Names  in  Zoology. 

Morum  (Oniscidia)  ninomiyai,  new  species 

Figs.  1-5 
Morum  sp.  Kaicher,  1983,  #3730,  3  figs.,  "off  Thailand". 
(Illustrations  of  the  paratype  of  this  species.) 

Diagnosis:  Member  of  the  Morum  (0.)  cancel- 
latum (Sowerby,  1824) -M.  (0.)  watanabei 
Kosuge,  1981,  complex.  Distinguished  by  having 
the  outer  lip  and  the  thin  parietal  shield  sculp- 
tured by  numerous,  fine,  thread-like  ridges  (Fig. 
5). 

Description:  Shell  medium  size  for  genus,  at- 
taining 40-1-  mm  in  height,  pyriform,  attenuated 
at  base.  Spire  low,  extended;  protoconch  erect 
and  papillate,  composed  of  IV2  glossy,  smooth 
whorls.  Postnuclear  whorls  5  in  number,  low 
conical,  weakly  shouldered  below  the  suture, 
first  and  second  whorls  weakly  cancellated,  sub- 
sequent whorls  strongly  cancellated,  with  14 
prominent  spinose  axial  ridges  per  whorl 
crossed  by  10  moderately  weak  spiral  cords  to 
form  a  blade-like,  hooked  spine  at  the  juncture 
of  the  spiral  and  axial  ribs;  spines  most  promi- 
nent at  the  shoulder.  Intervarical  areas  with  8  to 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      97 


FIGS.  .\-5.Morum  (Oniscidia)  ninomiyai,  new  species.  1  and  2,  Holotype,  AMNH  no.  221241.  3-5,  Paratype.  T.  Ninomiya col- 
lection (Fig.  5,  coated  to  show  sculptural  details;  photograph  courtesy  of  S.  D.  Kaieher).  6-9.  Morum  (Oniscidia)  cancellatum 
(Sowerby,  1821);  after  Emerson,  1985,  pi.  1,  figs.  15-18.  6  and  7,  off  Taiwan,  AMNH  no.  183783.  8  and  9,  Lectotype,  BM(NH) 
no.  197744,  "China  Seas"  (photographs  courtesy  of  A.  Beu).  All  figures  except  figure  5  approximately  x  1. 


10  evenly  spaced  axial  lamellae.  Aperture  nar- 
row, elongate,  semicrescentic  in  outline. 
Parietal  shield  moderately  large,  thin  with  outer 
edge  raised  and  covered  by  numerous,  fine,  ir- 
regular linear  lirations,  giving  the  appearance 
of  fine  threads.  Outer  lip  thickened,  crenulated 
and  weakly  toothed,  with  about  10  primary 
teeth  forming  inconspicuous  linear  projections 
on  the  inner  labial  margin,  2  secondary  teeth 
formed  between  each  pair  of  primary  teeth. 
Anal  sulcus  shallow;  siphonal  canal  short,  widely 
open,  weakly  recurved.  Operculum  not  known. 

Color:  Nucleus  shiny,  buff;  postnuclear  whorls 
with  whitish  base  color,  overlaid  with  flecks  of 
reddish  brown;  4  widely  spaced,  interrupted 
brownish  spiral  bands  on  body  whorl  (1  band  on 
shoulder,  2  on  either  side  of  midbody  area,  1 
near  the  base).  Aperture  milky  white;  outer  lip 
and  parietal  shield  white  and  surface  thinly 
glazed  (in  holotype). 

Measurements:  Holotype,  41.8  mm  in  height, 


25.7  mm  in  width;  paratype,  40.6  mm  in  height, 

25.8  mm  in  width. 

Type  locality:  off  Phuket  Island,  Thailand 
(8°N,  98.22°E),  Andaman  Sea,  dredged  in  50  to 
150  meters. 

Type  specimens:  holotype,  AMNH  221241 
(Figs.  1,  2)  from  type  locality;  paratype,  T. 
Ninomiya  Collection,  from  "off  Thailand, 
Andaman  Sea,  ex-Carl  Withrow  Collection, 
April  28,  1980",  (Figs.  3-5). 

Distribution:  Known  only  from  the  type  locali- 
ty and  adjacent  waters. 

Remarks:  The  new  species  superficially 
resembles  specimens  oi  Mo7iim  (0.)  cancellatum 
(Figs.  6,  7,  and  8,  9  (lectotype)  and  M.  (0.) 
watanabei  Kosuge  (see  Kosuge,  1981,  pi.  33, 
figs.  1,  2  holotype;  Emerson,  1985,  figs.  11-14), 
but  differs  in  the  number  of  axial  and  spiral 
ridges,  the  dentition  of  the  outer  lip  and  the 
unique  sculpture  of  the  parietal  shield  (Fig.  5). 


98      THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


Acknowledgments 

We  are  grateful  to  Taizo  Ninomiya  for 
depositing  the  holotype  of  this  new  taxon  in  the 
type  collection  of  moUusks  of  the  American 
Museum  of  Natural  History.  Richard  L. 
Goldberg  of  Fresh  Meadows,  New  York, 
William  Liltved  of  San  Francisco,  California, 
and  Peggy  Williams  of  Sarasota,  Florida  kindly 
provided  information  on  living  Morum  observed 
or  photographed  in  the  field  and  laboratory. 
Their  generous  cooperation  is  much  appre- 
ciated. Alan  G.  Beu  of  the  New  Zealand  Geologi- 
cal Survey,  and  Sally  Diana  Kaicher  of  St. 
Petersburg,  Florida,  generously  provided  the 
photographs  for  figures  8,  9  and  5,  respectively. 

I  am  also  grateful  to  Dr.  Roger  N.  Hughes  of 
the  University  College  of  North  Wales  for  his 
kindness  in  keeping  me  informed  of  his  anatom- 
ical studies  of  Morum. 

I  thank  my  AMNH  colleagues  for  their  con- 
tributions to  this  study:  Walter  E.  Sage  HI  for 
technical  assistance,  Stephanie  Crooms  for 
word-processing  the  manuscript,  and  Stephen 
Butler  for  the  photography,  except  as  noted. 

LITERATURE  CITED 

Adams,  Arthur  1855.  Descriptions  of  new  genera  and 
species  of  gasteropodus  |.s-(c"]  Mollusca.  Pror.  Zool.  Soc. 
London  for  1853,  21(259):  182-186  (May  16,  1855). 


Boss,  K.  J.  1982.  Mollusca,  in  S.  P.  Parker,  ed.,  Synopsis 
and  Classification  of  Living  Organisms,  1  McGraw-Hill 
Book  Co.,  New  York,  pp.  945-1166,  illus. 

Emerson,  W.  K.  1981.  Two  new  Indo-Pacific  species  of 
Morum  (Gastropoda:  Tonnacea).  The  Nautilus  95(3): 
101-105,  7  figs, 

1985.  Remarks  on  some  western  Pacific  species 

of  Morum  (Gastropoda:  Tonnacea).,  in  J.  M.  Lindsay,  ed.. 
Stratigraphy,  palaeontology,  malacology  papers  in  honour 
of  Dr.  Nell  Ludbrook.  Spec.  Publ.  So.  Aust.  Dept.  Mines 
and  Energy  5,  pp.  51-56,  2  pis. 

Hughes,  R.  N.  1986.  Anatomy  of  the  foregut  of  Morum 
Roding,  1798  (Gastropoda:  Tonnacea)  and  the  taxonomic 
misplacement  of  the  genus.  The  Veliger  29(1),  in  press. 

Kaicher,  S.  D.  1983.  Card  Catalogue  of  World-Wide  Shells, 
Pack  #36,  Cassidae  &  Oocorythidae,  card  #3730. 

Kosuge,  Sadao  1981.  Descriptions  of  two  new  species  of  the 
genus  Morum  with  remarks  on  the  Recent  species  from 
Philippines.  Bull.  Inst.  Malac.  Tokyo  1(7):101-104,  pi.  33. 

Kuroda,  Tokubei  and  Tadashige  Habe,  in  T.  Habe.  1961. 
Colored  Illustrations  of  the  Shells  of  Japan,  2,  Hoikusha, 
Osaka,  Appendix,  p.  1-41. 

Morch,  0.  A.  L.  1852.  Catalogus  conchyliorum  .  .  .  de  Yoldi., 
Regis  Daniae.  Copenhagen.  Fac.  1.  170  p. 

Raven,  J.  G.  M.  1985.  Homonymy  in  the  families  Harpidae 
Hawle  &  Corda,  1847  (Trilobita)  and  Harpidae  Bronn, 
1849  (Mollusca,  Gastropoda).  Z.N.(S.)  2331.  Bull.  Zool. 
Nom.  42(l):79-80. 

Rehder,  H.  A,  1973.  The  family  Harpidae  of  the  world.  Indo- 
Pacific  Mollusca  3(16):207-274,  pis.  183-247. 

Sowerby,  G.  B.  I.  1824.  The  Genera,  of  Recent  and  Fossil 
Shells  1,  Oniscia,  pi.  233. 

Thiele,  Johannes.  1929.  Handbuch  der  Systematischen 
Weichtierkunde.  Jena.  Bd  1,  teil  1.  pp.  1-376,  470  text 
figs. 


A  NEW  DEEP-WATER  SPECIES  OF  LEPIDOPLEURUS 
(POLYPLACOPHORA)  FROM  THE  VENEZUELA  BASIN. 

Antonio  J.  Ferreira^ 

Research  Associate,  Department  of  Invertebrate  Zoology 

California  Academy  of  Sciences 

San  Francisco,  CA  94118 

ABSTRACT 
A  new  species  o/Lepidopleurus /rom  the  depths  of  the  Venezuela  Basin,  Carib- 
bean Sea,  is  described  and  compared  with  L.  scrippsianus  Ferreira.  1980,  from 
the  Eastern  Pacific. 


From  October  to  December  1981,  the  Naval 
Ocean  Research  and  Development  Activity 
(NORDA),  NSTL  Station,   MS,  conducted  in- 


'Published  posthumously.  Died  1986. 


tense  physical  and  biological  investigations  of 
deep-sea  sediments  in  the  Venezuela  Basin  from 
aboard  USNS  Bartlett  (cruise  1301-82).  Among 
the  benthic  macrofauna  were  6  specimens  of 
chitons:  3  specimens,  ca.  13,  11,  and  9  mm  long 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS      99 


3  4  5 

FIGS.  1-5,  Lepidopleurus  bartletti  Ferreira,  spec  nov.:  Holotype  (USNM  859000),  side 
view.  2,  view  of  posterior  valve.  3,  first  and  second  valve.  4,  paratype,  posterior  valve.  5, 
paratype,  under  view,  valves  i,  iv  and  viii. 


(sta.  40:  15°08'N,  69°  12^,  depth  3867  to  4009 
m,  28  Oct.  1981);  1  specimen,  ca.  15  mm  long 
(sta.  63:  13°45'N,  67°45'W,  depth  5046  m,  8 
Nov.  1981);  1  specimen,  ca.  10  mm  long  (sta.  65: 
13°45'N,  67°45'W,  depth  5046  m,  9  Nov.  1981); 
and  1  specimen,  1.3  mm  long  (sta.  88:  13°30'N, 
64°45'W,  depth  3516  to  3550  m,  25-26  Nov. 
1981).  All  chitons  were  found  on  pieces  of  wood 
brought  up  in  trawls.  Study  of  this  material, 
generously  made  available  through  Michael  D. 
Richardson  and  Paula  M.  Mikkelsen,  revealed  a 
new  species  here  allocated  to  the  genus  Lepido- 
pleurus Risso,  1826. 

Order  Neoloricata  Berhenhayn,  1955 

Family  Lepidopleuridae  Pilsbry,  1892 

Genus  Lepidopleurus  Risso,  1826 

Lepidopleurus  bartletti  Ferreira,  spec.  nov. 

Fibres  1-6 

Diagnosis:  Small  (up  to  15  mm  long)  white 

chitons.    Valves    somewhat    rugose;    posterior 

edges  moderately  angled;  surface  covered  with 

microgranular  cuticle  over  sculptureless,  chalky 


tegmentum.  End  valves  remarkably  flat;  mucro 
slightly  anterior;  lateral  areas  not  prominent, 
with  some  concentric  growth  rugae  continuing 
into  central  areas.  Gills  posterior.  Girdle  thick; 
upper  surface  carpeted  with  small  spiculoid 
elements  with  occasional,  thin,  straight,  long 
spicules  interspersed;  undersurface  reduced  to 
fine  cuticle  devoid  of  scales  or  spicules.  Radula 
median  teeth  wider  in  back  than  in  front,  with 
tricuspid  major  lateral  teeth  and  rakelike 
spatulate  teeth. 

Type  material:  Holotype  (USNM  859000)  and 
paratypes  (CAS  059845;  IRCZM  061:085; 
LACM  2123). 

Other  material:  Specimen  1.3  x  0.9  mm,  from 
sta.  88,  tentatively  identified  as  L.  bartletti 
(CAS  059846)  but  not  given  paratypic  status  in 
view  of  its  small  size  and  ill-defined,  juvenile 
characters. 

Type  locality:  Venezuela  Basin,  Caribbean 
Sea,  13°45'N,  67 "45^,  at  depth  of  5046  m. 
(Station  63,  roughly  150  mi.  N.E.  of  Bonaire, 
Netherlands  Antilles). 


100    THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


Description:  Holotype  (from  USNV  Bartlett 
sta.  63),  preserved  in  alcohol,  curled,  ca.  15  mm 
long,  10  mm  wide  (figs.  1-3).  Valves  subcarinate, 
moderately  beaked,  with  angled  posterior 
edges.  Shell  surface  covered  with  microgranular 
cuticle,  easily  scraped  off  to  show  chalky  white, 
sculptureless  tegmentum  beneath.  Anterior 
valve  remarkably  flat.  Lateral  areas  of  inter- 
mediate valves  not  prominent  except  for 
greater  valve  thickness  and  concentric  growth 
rugae  often  continuing  unto  sculptureless  cen- 
tral areas.  Posterior  valve  extremely  flat;  macro 
slightly  anterior,  poorly  defined.  Gills  posterior, 
about  20  plumes  per  side,  extending  40%  of 
length  of  foot.  Girdle  creamy  white,  thick,  with 
conspicuous  straight,  thin,  glassy  spicules  inter- 
spersed on  background  of  much  smaller,  thin- 
ner, closely  packed  spiculoid  elements. 

Paratypes  (from  USNV  Bartlett  sta.  40  and 
sta.  63.)  very  similar  to  holotype.  Paratype  from 
sta.  40,  13  mm  long,  disarticulated:  Cuticle  easi- 
ly removable  from  valves  uncovering  chalky, 
sculptureless  tegmental  surface  beneath  (fig.  4). 
Articulamentum  white  with  no  insertion  teeth 
or  slits  (fig.  5).  Valve  i,  5.2  mm  wide,  3.2  mm 
long,  1.2  mm  high  at  posterior  edge.  Valve  v,  6.0 
mm  wide,  2.0  mm  long  in  midline.  Valve  viii,  5.1 
mm  wide,  3.7  mm  long,  1.0  mm  high  at  anterior 
edge,  slightly  convex  in  premucro  area,  slightly 
concave   in   postmucro  area.   Sutural  laminae 


short,  triangular;  sinus  very  wide;  on  valve  viii, 
relative  width  of  sinus  (width  of  sutural  sinus/ 
width  of  sutural  lamina)  is  1.8.  Girdle  dorsal  sur- 
face densely  carpeted  with  blunt-ended,  cigar- 
shaped,  spiculoid  elements  up  to  100  x  25  nm 
(up  to  120  X  35  ^im  at  sutures),  often  longi- 
tudinally striated  (fig.  6-A),  interspersed 
straight,  vaguely  striated,  glassy  spicules  up  to 
300  X  28  ^m  (fig.  6-B),  interspersed;  undersur- 
face  reduced  to  fine  cuticle  without  scales, 
spicules  or  other  elements;  girdle  bridges  emp- 
ty. Radula  6  mm  long,  comprising  55  rows  of 
mature  teeth;  median  teeth  (fig.  6-A)  80  ^im 
long,  30  ^m  wide  at  anterior  blade,  enlarging 
posteriorly  to  60  ^m;  first  lateral  teeth  about  80 
^<m  long,  with  socketlike  concavity  at  anterior 
end;  major  lateral  teeth  with  tricuspid  head  (fig. 
6-B)  about  70  jum  wide,  100  jim  long  at  middle 
cusp;  spatulate  teeth  150  ^m  wide  with  fascicu- 
late, rakelike  appearance  (fig.  6-C);  outer  mar- 
ginal teeth  90  ^m  long,  120  ^m  wide  (width/ 
length,  0.75). 

Distribution:  Lepidopleurus  bartletti  is  known 
only  from  the  general  area  of  the  type  locality  in 
the  central  lower  Caribbean  at  great  depths. 

Remarks:  In  addition  to  L.  bartletti,  only  two 
other  living  species  oi Lepidopleurus  are  known: 
L.  cajetanus  (Poli,  1791),  a  common  intertidal  to 
sublitoral  European  species,  and  L.  scrippsia- 
nus  Ferreira,  1980,  collected  at  over  2500  m  in 


FIG.  6.  Lepidopleurux  bartletti  Ferreira,  spec,  nov.:  Paratype  (CAS  0.')984.5),  radular  -  (A)  median  and  first  lateral  teeth; 
(B)  head  of  major  lateral  tooth;  (C)  head  of  spatulate  tooth;  (U)  interspersed  spicule;  (E)  background  spiculoid  elements. 
Scale  bar  100  nm. 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS    101 


the  eastern  Pacific,  SW  of  Cabo  San  Lucas,  Baja 
California,  Mexico.  A  fossil  species,  L.  rnoro- 
zakierisis  Itoigawa,  Nishimoto  and  Tomida, 
1977,  is  known  from  the  Miocene  Morozaki 
group  of  central  Japan. 

Lepidopleurus  bartletti  is  quite  similar  to  L. 
scrippsiayms  from  which  it  differs  in  1)  more 
delicate,  less  rugose  valves,  2)  posterior  edge  of 
valves  forming  much  wider  angle,  3)  lateral 
areas  much  less  accentuated,  4)  extremely  flat 
end  valves,  5)  anterior  mucro,  and  6)  valve 
covering  cuticle  [not  seen  in  specimens  of  L. 
scrippsianus],  giving  bartelli's  valves  a 
microgranular  appearance  which  contrasts  with 
the  sculptureless.  chalky  tegmental  surface 
beneath.  Notable  features  common  to  L.  bart- 
letti and  L.  scrippsianus  are  1)  girdle  elements, 
2)  girdle  undersurface  reduced  to  cuticle 
without  spicules  or  scales,  and  3)  radula  with 
unusually  shaped  median  teeth  and  rake-like 
spatulate  teeth. 

The  species  is  named  bartletti  in  honor  of  the 
USNS  Bartlett  for  her  central  role  in  this  study 
of  the  Venezuela  Basin. 

Acknowledgments 

I  express  my  appreciation  to  Paula  M.  Mikkel- 


sen,  Indian  River  Coastal  Zone  Museum,  Harbor 
Branch  Foundation,  Inc.,  Fort  Pierce,  Florida, 
who  entrusted  these  specimens  to  my  care  and 
study.  William  G.  Lyons  of  Florida's  Depart- 
ment of  Natural  Resources  very  kindly  made 
many  improvements  to  the  manuscript. 

LITERATURE  CITED 

Bergenhayn,  J.  R.  M.  1955.  Die  fossilen  schwedischen 
Loricaten  nebst  einer  vorlaufigen  Revision  des  systems 
der  ganzen  Klasse  Loricata.  Lunds  Univ.  Arsskrift. 
(Avd.2,  N.S.)  51  (8):l-43,  2  pits.  [Kungl.  Fysiogr.  Sallsk. 
Hand!.  N.F.  66  (8):3-42,  2  tables]. 

Ferreira,  A.  .J.  1980.  A  new  species  of  Lepidopleurus  Risso, 
1826  (Mollusca:  Polyplacophora)  in  the  deep  waters  of  the 
eastern  Pacific.  The  VeUger  23(1):55-61,  5  text  figs.,  1  pi. 

Gray,  J.  E.  1821.  A  natural  arrangement  of  IVIollusca, 
according  to  their  internal  structure.  London  Medic. 
Repos.  15:229-239. 

Itoigawa,  J.,  H.  Nishimoto  and  S.  Tomida.  1977.  Lepido- 
pleurus rtwrozakiensis,  a  new  fossil  Polyplacophora 
from  the  Miocene  Morozaki  group,  central  Japan.  Bull. 
Mizunami  FossU  Mus.  4:55-59,  2  text  figs.,  pis.  14-15. 

Pilsbry.  H.  A.  1892.  Polyplacophora.  Manual  of  Conchology 
14:1-64;  pits.  1-15. 

Poli,  I.  X.  1791-1827.  Testacea  utrisque  Siciliae  eorumque 
historia  et  anatome  tabulis  aeneis  illustrata.  Vol.  1  (1791). 

Risso,  A.  1826.  Histoire  naturelle  des  principales  produc- 
tions de  I'Europe  Meridionale  et  particulierement  de 
celles  des  environs  de  Nice  et  des  Alpes  maritimes.  Vol. 
4,  439  pp.,  12  pits.,  183  figs.  Paris  &  Strasbourg:  F.-G. 
Levrault.  5  vols. 


A  NEW  SPECIES  OF  LYRIA  (GASTROPODA:  VOLUTIDAE) 
FROM  THE  ARABIAN  SEA 

William  K.  Emerson  and  Walter  E.  Sage  III 

Department  of  Invertebrates 

American  Museum  of  Natural  History 

New  York,  NY  10024 

ABSTRACT 
Lyria  leslieboschae,  new  species,  is  described  from  off  Masirah  Island, 
Sultanate  of  Oman,  Arabian  Sea.  Known  only  from  a  few  beach  specimens,  this 
new  taxon  is  distingwished  by  discrete  differences  in  shell  morphology  from  the 
superficially  similar  species,  Lyria  lyraeformis  (Swainson,  1821),  which  occurs 
off  the  coast  of  Kenya,  East  Africa. 


Donald  and  Eloise  Bosch  resided  for  nearly 
thirty  years  in  Oman,  where  she  was  a  teacher 
and  he  was  a  practicing  physician.  During  this 
time,  they  became  knowledgable  shell  collectors 


and  eventually  co-authored  an  illustrated  guide, 
"Seashells  of  Oman",  which  was  published  in 
1983.  Among  specimens  they  recently  submit- 
ted to  us  for  study  were  four  examples  of  a  new 


102    THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


FIGS.  1,  3-5.   Hulutype  of  Lyria  kslieboschae  new  species,  AMNH  no.  221B5.',.  3-5,  dorsal,  apertural  and  oblique  views,  x  1.  1, 
detail  of  early  whorls,  x  2. 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS    103 


FIGS.  2,  6-8,  Lyria  lyraeformis  (Swainson.  1821) 
oblique  views,  x  1.  2,  detail  of  early  whorls,  x2. 


trawled  off  the  coast  of  Kenya,  AMNH  21358.3.  6-8,  dorsal,  apertural,  and 


species  of  Lyria  found  for  the  first  time  in 
January,  1985,  at  Masirah  Island.  We  are  ex- 
tremely pleased  to  name  this  exquisite  volute  in 
honor  of  their  daughter-in-law,  Mrs.  Leslie 
Bosch,  who  actively  participated,  together  with 
her  husband,  David,  on  ten  annual  collecting  ex- 
peditions of  the  Bosch  family  to  this  locality. 

Lyria  leslieboschae  new  species 

Figs.  1,  3-.5 

Diagnosis:  In  outline  and  general  appearance 
the  shell  resembles  Lyria  lyraeformis  (Swain- 
son, 1821,  pi.  54,  2  figs.;  Weaver  and  duPont, 
1970,  pi.  5  A,  B;  Abbott  and  Dance,  1982,  p.  213, 


1  fig.;  and  Okutani,  1983,  pi.  31,  fig.  8;  here  illus- 
trated, figs.  2,  6-8),  but  differs  in  possessing  a 
blunt,  bulbous  nucleus,  which  lacks  a  calcarella 
(cf.  figs.  1,  2),  an  anal  canal  that  is  narrow,  open 
and  flares  posteriorly  to  the  region  just  below 
the  subsutural  spiral  band  (cf.  figs.  4,  7),  as  well 
as  by  axial  sculpture  that  is  less  strongly 
developed  on  the  body  whorl,  and  a  distinctive 
color  pattern  that  is  more  diffused. 

Description  of  holotype,  figures  1,  3-5:  Shell 
large  for  genus,  attaining  130  mm  in  height, 
elongate-fusiform,  spire  high,  attenuated. 
Nucleus  of  2V2  smooth  whorls,  without  a  cal- 
carella. Teleoconch  of  7V2  rounded  whorls,  in- 


104    THE  NAUTILUS 


July  30,  1986 


Vol.  100  (3) 


dented  at  the  suture;  axial  sculpture  weakly 
developed  on  body  whorl,  numbering  21  on 
penultimate  whorl,  more  evenly  defined  on  the 
early  whorls  than  on  the  penultimate  and  body 
whorls.  Spiral  sculpture  best  expressed  on  the 
wide,  spirally  indented  subsutural  bands,  each 
with  6  spiral  lirae  acutely  raised  at  the  points  of 
alignment  with  the  axial  ribs.  Aperture  narrow, 
gaping  anteriorly,  with  a  short  wide  siphonal 
canal,  terminating  posteriorly  in  a  long,  narrow, 
open  anal  canal,  near  the  base  of  the  spiral  band 
below  the  suture.  Columella  with  three  anterior 
plaits;  parietal  region  without  plaits.  Exterior 
surface  above  the  siphonal  canal  with  7  spiral 
ridges. 

Base  color  a  cream-tan,  overlaid  with  inter- 
rupted bands  of  reddish  brown  formed  in  the 
subsutural  areas  and  on  the  medial  and  basal 
regions  of  the  body  whorl.  Reddish  brown  spiral 
lines  on  the  axial  ribs.  Reddish  brown  irregular 
axial  lines  on  the  nucleus.  Interior  of  aperture 
and  columella  glossy,  colored  a  tannish  cream, 
with  a  bluish  tint.  The  coloration  of  live-taken 
specimens  can  be  expected  to  be  more  vividly 
expressed  than  in  this  well-preserved  beach 
specimen,  which  had  several  large  colonies  of 
bryozoa  encrusting  the  interior  of  the  aperture 
at  the  time  of  collection. 

Nothing  is  known  of  the  radula  or  soft  parts. 

Measurements:  Holotype,  AMNH  no.  221655, 
129.3  mm  in  height,  38.4  mm  in  width.  Para- 
types,  AMNH  no.  221656,  Paratype  A,  85.5  mm 
in  height,  31.3  mm  in  width,  Paratype  B,  87.9 
mm  in  height,  31.9  mm  in  width,  Paratype  C, 
91.8  mm  in  height,  29.4  mm  in  width.  (The 
paratypes  lack  complete  spires). 

Type  locality:  About  8  km  south  of  the  village 
of  Haql,  on  the  southeastern  coast  of  Masirah 


Island,  Oman. 

Distribution:  Known  only  from  the  tj^De 
locality. 

Remarks:  The  only  known  specimens  were 
washed  ashore  by  the  heavy  surf  on  the  Indian 
Ocean  side  of  the  island.  On  this  exposed  beach, 
Dr.  Bosch  {in  litteris,  May  9,  1986)  also  reported 
finding  dead  specimens  of  Festilyria  festiva 
(Lamarck,  1811)  and  Conus  lischkeanus  cf. 
tropicensis  Coomans  and  Filmer,  1985,  species 
which  are  known  to  occur  in  moderately  deep 
offshore  waters.  The  present  species  apparently 
lives  at  moderate  depths  and  will  require 
SCUBA  diving  and/or  dredging  operations  to 
collect  living  specimens. 

Acknowledgments 

We  thank  Dr.  and  Mrs.  Bosch  for  their  kind- 
ness in  calling  to  our  attention  this  most  elegant 
addition  to  the  Volutidae  and  for  depositing  the 
typological  specimens  in  the  type  collection  of 
mollusks  of  the  American  Museum  of  Natural 
History.  We  also  thank  Stephanie  Crooms  for 
word-processing  the  manuscript  and  Stephen 
Butler  for  the  photography. 

LITERATURE  CITED 

Abbott,  R.  T.  and  Dance,  S.  P.  1982.  Compendium  Of  Sea- 
shells.  E.  P.  Dutton,  New  York,  411  pp.,  illus.  in  color. 

Bosch,  D.  and  E,  1982.  Seashells  Of  Oman.  Longman  Group 
Ltd.,  London  and  New  York,  206  pp.,  illus.  in  color. 

Okutani,  T.  1983.  World  Seashells  of  Rarity  and  Beauty. 
Kawamura  Collection.  National  Science  Museum,  Tokyo, 
i-iii  +  1-12  pp.,  48  pis.  in  color. 

Swainson,  W.  1821.  Zoolugical  Illustrations,  London,  1, 
pi.  54. 

Weaver,  C.  S.  and  duPont,  J.  E.  1970.  Living  Volutes:  a 
monograph  of  Recent  Volutidae  of  the  world.  Delaware 
Mus.  Nat.  Hist.  Mong.  Ser.,  No.  1.  xv  +  374  pp.,  79  col. 
pis.,  44  figs.,  13  maps. 


BOOK  REVIEW 


Hong  Kong  Seashells  by  John   D.   Orr.   1985. 

Urban  Council,  Hong  Kong.  122  pp.,  94  color 

photos.  Paperback. 

The  author  presents  60  beautiful,  close-up 
photographs  of  living  specimens  of  cowries, 
olives,  cones,  Stromhus  conchs  and  helmet 
shells,  as  well  as  the  shells  of  84  other  common, 
shallow  water  bivalves  and  gastropods  of  the 


Hong  Kong  area.  Hints  on  collecting,  classifying 
and  studying  mollusks  are  included.  The  spec- 
tacular photographs  of  living  specimens  make 
up  for  the  few  included  species,  lack  of  authors 
and  dates,  and  occasional  typographical  errors. 
Cymatium  clandestinum  on  page  102  is  really  a 
worn  Cantharus  (Pollia)  probably  undosa 
(Linnaeus,  1758). -/2.  T.  Abbott. 


Vol.  100  (3) 


July  30,  1986 


THE  NAUTILUS    105 


LITHARCA  LITHODOMUS  AND  ADAPTIVE  RADIATION 
IN  ARCACEAN  PELECYPODS 


David  Nicol 

Box  14376 

University  Station 

Gainesville,  FL  32604 


and 


Douglas  S.  Jones 

Florida  State  Museum 
Gainesville,  FL  32611 


ABSTRACT 

Litharca  lithodomus  is  a  rock-boring  arcid  (subfamily  Litharcinae  Frizzell, 
19Jf6)  that  has  a  geographic  range  from  Nicaragua,  to  Peru.  It  is  not  known  as  a 
fossil,  and  this  species  may  be  less  than  one  million  years  old.  Litharca 
lithodomus  is  most  likely  a  mechanical  rock-borer.  The  possible  ancestral  species 
is  Barbatia  (Cucullaearca)  reeveana,  which  is  commonly  found  throughout  the 
present  range  of  Litharca  lithodomus.  Barbatia  (Cucullaearca)  reeveana  nestles 
in  crevices  in  rocks  and  its  shape  is  frequently  distorted.  The  change  of  habit  from 
nestling  to  rock-boring  is  not  great,  and  this  change  is  seen  in  some  other  species 
in  dijferent  families  of  pelecypods. 

The  earliest  arcaceans,  which  appeared  during  the  Middle  Devonian,  were 
epibyssate.  Shallow-burrowing  species  of  arcaceans  probably  did  not  occur  before 
the  Triassic,  and  nestling  species  probably  appeared  no  later  than  the  Paleocene. 
Fresh-water  and  rock-boring  arcaceans  seem  to  be  confined  to  the  Recent. 


Many  large  families  or  superfamilies  of  sus- 
pension-feeding pelecypods  exhibit  more  adap- 
tive radiation  than  all  of  the  Protobranchia, 
which  are  deposit  feeders.  One  of  these  large 
superfamilies,  the  Arcacea,  is  diverse  and  wide- 
ly distributed  in  the  seas  today,  ranging  from 
the  tropics  to  the  cold  water  of  the  Antarctic,  as 
well  as  to  abyssal  and  hadal  regions. 

The  earliest  true  arcaceans  appear  suddenly 
in  the  Middle  Devonian  (Givetian  Stage)  about 
380  million  years  ago.  Most  paleontologists  have 
said  that  the  Cyrtodontidae  represent  the 
ancestral  stock  that  gave  rise  to  the  arcaceans, 
based  on  the  similarity  of  the  hinge  teeth  and 
ligament  in  some  members.  A  few  have  doubted 
that  the  cyrtodonts  were  the  ancestors  of  the 
arcaceans  (Nicol,  1984).  Perhaps  a  more  likely 
group  to  have  given  rise  to  the  arcaceans  was 
the  Ambonychiidae.  Some  ambonychiids  have 
ligaments  and  hinge  teeth  similar  to  the 
arcaceans  and  many  were  byssally  attached 
whereas  the  Cyrtodontidae  were  burrowers  ac- 
cording to  Stanley  (1972).  Furthermore,  while 
the  Ambonychiidae  were  common  in  the  Middle 
Devonian,  the  Cyrtodontidae  were  nearly  ex- 
tinct by  that  time.  It  is  also  easier  to  derive  the 
byssally  attached  arcacean  Parallelodon  from  a 


byssally  attached  ambonychiid  rather  than  a 
burrowing  cyrtodontid. 

It  is  generally  agreed  that  the  first  arcaceans 
were  byssally  attached  to  the  substrate,  and 
Parallelodon  has  a  rectangular  valve  outline 
that  resembles  Area,  sensu  stricto.  Some  of  the 
Paleozoic  arcaceans  were  probably  endobyssate, 
and  this  life  habit  occurred  in  some  species  of 
Parallelodon.  However,  arcaceans  are  not  com- 
mon and  diverse  throughout  the  Paleozoic,  and 
the  great  adaptive  radiation  of  this  superfamily 
began  in  the  Jurassic  and  continued  throughout 
the  Cretaceous  (Nicol,  1984). 

The  first  burrowing  arcacean  may  have  been 
Macrodontella,  which  occurs  in  the  Triassic  (text 
fig.  1).  These  early  burrowing  arcaceans  did  not 
appear  until  at  least  150,000,000  years  after  the 
first  arcaceans  appeared.  Other  burrowing  arca- 
ceans occur  amongst  the  cucullaeids  in  the 
Jurassic  and  Cretaceous,  and  the  glycymeridids 
and  Trigonarca  during  the  Cretaceous.  Some  of 
the  Cretaceous  limopsids  were  probably  bur- 
rowers  but  some  were  endobyssate  (Oliver, 
1981).  Epibyssate,  endobyssate,  and  burrowing 
arcaceans  continue  to  the  Recent  with  burrow- 
ing species  probably  outnumbering  the  epibys- 
sate and  endobyssate  forms. 


106    THE  NAUTILUS 


July  30,  1986 


GEOLOGIC 
AGE 


ROCK-BORING  FRESHWATER 

BYSSATE  NESTLING 


Vol.  100(3) 
FREE-BURROWING 


RECENT 

PALEOCENE 
(65  mya) 


TRIASSIC 
(230  mya) 


LItharca 


DEVONIAN 
(380  mya) 


•  Scaphula 


Acar 


Macrodontella 


Parallelodon 

I 
I 
I 

Ambonychlidae    ? 


FIG.  1 .  Diagram  showing  adaptive  radiation  in  the  Arcacea. 


Some  species  of  Acar  are  true  nestlers,  and 
this  genus  appeared  in  the  Paleocene,  about 
65,000,000  years  ago.  Scaphula  is  an  arcacean 
that  has  invaded  brackish  and  fresh  water  dur- 
ing the  Recent. 

The  most  unusual  adaptation  of  the  arcaceans 
occurs  in  Litharca  lithodom.ics  (Sowerby,  1833), 
a  rock-borer  (plate  1).  The  genus  is  monotypic 
and  is  known  only  from  the  Recent.  Keen  (1971) 
stated  that  the  range  of  the  species  extends 
from  Nicaragua  to  Peru,  and  it  appears  to  be 
most  common  along  the  coast  of  Ecuador. 
Olsson  (1961)  claimed  that  Litharca  is  so  com- 
mon in  some  areas  of  Ecuador  that  fishermen 
use  it  for  food  and  bait.  Litharca  lithodomus  is  a 
relatively  large  arcid,  and  Frizzell  (1946)  had 
one  specimen  that  was  91.1  mm  in  length. 
Olsson  (1961)  stated  that  the  adult  shell  reaches 
a  length  of  nearly  100  mm. 

The  shell  of  Litharca  is  unusually  thick  for  a 
rock-borer,  and  only  some  species  of  rock-boring 
tridacnids  have  slightly  thicker  shells.  In  other 
words,  rock-boring  pelecypods  commonly  have 
very  thin  shells  as  seen  in  the  mytilids,  pholads, 
petricolids,  and  myids.  A  thin  shell  may  have  an 


advantage  in  that  it  can  be  more  easily  and  fre- 
quently moved  within  the  borehole.  Further- 
more, a  thick  shell,  as  in  Mercenaria,  is  not 
needed  for  protection  from  carnivorous  gastro- 
pods and  crustaceans.  Shells  of  mechanical  rock- 
boring  pelecypods  commonly  consist  of  arago- 
nite,  but  the  nacreous  structure  found  in 
unionids  and  Pteriidae  is  absent.  Nacreous 
shells  are  friable  and  would  not  be  suitable  for 
mechanical  rock  boring. 

One  of  the  most  unusual  features  of  Litharca 
lithodomus  is  the  greatly  elongated  anterior  end 
(Plate  1).  The  ligament  is  wholly  prosodetic  and 
lacks  a  posterior  limb  (i.e.,  the  ligament  lacks 
chevrons).  This  lack  of  a  posterior  limb  of  the 
ligament  is  also  seen  in  the  Cretaceous 
glycymeridid  Postligata  (Nicol  and  Jones,  1984). 
Frizzell  (1946)  stated  that  the  beaks  are  slightly 
opisthogyrate,  but  Olsson  (1961)  said  that  they 
are  probably  prosogyrate.  The  beaks  are  most 
likely  opisthogyrate  because  of  the  prosodetic 
ligament  and  the  extreme  elongation  of  the 
anterior  end  of  the  shell.  Prosodetic  ligaments 
and  opisthogyrate  beaks  are  quite  common  in 
glycymeridids.   A   byssus   is   present  in   adult 


Vol.  100  (3) 


July  30,  1986 


THE  NAUTILUS    107 


055l>)0 


PLATE  1.  A-D.  Litharca  lithodomus  (Sowerby).  Manta,  Ecuador.  A,  Exterior,  right  valve.  B,  Interior,  left  valve.  C,  Ventral 
view,  both  valves  showing  small  byssal  gape  (shell  length  86.5  mm).  D,  Dorsal  view,  both  valves  showing  truncated  posterior  to 
the  left.  A-D,  California  Academy  Sciences  #055190.  E-F,  Barhatia  iCucuUaearca)  reeveana  (Orbigny).  Perlas,  Panama.  E, 
Left  valve,  exterior.  F,  Left  valve,  interior  (shell  length  79.7  mm).  E-F.  Florida  State  Museum  #23624. 


Litharca  lithodomus,  but  the  byssal  gape  is  nar- 
row and  elongate,  and  the  byssus  is  flexible  so 
that  the  animal  is  able  to  move  downward  in  its 
borehole  (Olsson,  1961).  As  is  shown  in  Frizzell's 
figure  5  (1946),  the  valves  are  wedge-shaped 
with  the  most  convex  portion  near  the  posterior 
end.  The  anterior  end  is  narrow.  This  is  an 
unusual  shape  for  a  rock-borer. 

Thomas  (1974,  1976)  has  assumed  that  the 
boring  is  accomplished  by  chemical  means  in 
Litharca.  but  this  may  not  be  so.  Both  Frizzell 
(1946)  and  Olsson  (1961)  have  observed  that 
Litharca  bores  into  massive  sandstones  in 
Ecuador.  Are  these  sandstones  calcareous  or 
cemented  with  calcium  carbonate?  Field  and 
laboratory  observations  on  these  sandstones 
would  be  essential  to  come  to  a  final  conclusion 
as  to  the  mode  of  boring.  Just  because  Litharca 
has  a  weak  ligament  and  a  pitted  shell  exterior 
does  not  necessarily  rule  out  the  possibility  that 
it  may  bore  mechanically.  Furthermore,  of  the 
remaining  rock-boring  species  of  pelecypods. 


only  some  species  of  mytilids  bore  by  chemical 
means  whereas  other  species  of  rock-boring 
mytilids  are  mechanical  borers  (Yonge,  1951). 
Perhaps,  as  Thomas  implies  (1976),  both 
mechanical  and  chemical  means  of  boring  may 
be  employed  by  L  itharca. 

The  fact  that  at  least  one  arcid  species  has 
developed  a  rock-boring  habit  is  apparently  not 
as  incongruous  as  it  would  at  first  seem.  The 
two  most  unusual  features  found  in  Litharca 
lithodomus  that  are  generally  not  seen  in  other 
rock-boring  species  are  the  posterior  displace- 
ment of  the  beaks  and  the  hatchet-shaped  con- 
vexity of  the  valves  with  the  most  convex  region 
near  the  posterior  end.  Most  rock-borers  are 
cylindrical  in  shape,  and  the  beaks  are  either 
near  the  center  of  the  valves  or  near  the 
anterior  end.  The  byssus  is  not  unique  because 
rock-boring  mytilids  and  tridacnids  have  this 
organ  developed  in  the  mature  rock-boring 
stage  (Yonge,  1951).  Compared  to  some  other 
rock-boring  pelecypods,  Litharca  is  relatively 


108    THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


unspecialized. 

Rock-boring  pelecypods  are  most  diverse  in 
the  tropical  marine  regions.  They  do  not  occur 
in  the  abyssal  and  hadal  regions  of  the  oceans 
nor  in  fresh  water.  There  are  at  least  eight 
families  that  have  been  purported  to  have  one  or 
more  rock-boring  species  and  they  are:  Arcidae, 
Mytilidae,  Tridacnidae,  Petricolidae,  Hiatelli- 
dae,  Myidae,  Gastrochaenidae,  and  Pholadidae. 
The  Mytilidae  and  Pholadidae  comprise  most  of 
the  rock-boring  species,  but  there  are  probably 
no  more  than  100  living  species  of  rock-boring 
pelecypods.  Rock-boring  species  of  pelecypods 
are  of  average  to  large  size.  The  largest  rock- 
borers  are  three  species  of  Tridacna,  but  they 
are  the  smallest-sized  species  in  the  family  and 
none  attains  a  maximum  size  of  more  than  about 
400  mm.  Furthermore,  the  species  that  bores 
the  deepest  into  the  rock  is  Tridacna 
(Chametrachea)  crocea  Lamarck,  and  it  is  the 
smallest  of  the  three  species,  attaining  a  max- 
imum size  of  only  150  mm  (Rosewater,  1965).  It 
also  appears  that  species  of  pelecypods  that 
attain  a  size  of  less  than  20  mm  are  not  rock 
borers,  and  this  may  be  caused  by  a  lack  of  bor- 
ing capabilities  in  small-sized  species. 

It  was  assumed  by  Olsson  (1961)  and  Thomas 
(1978)  that  the  ancestor  of  Litharca  was  a 
species  of  Area.  An  hypothesis  ascribed  by 
Dechaseaux  (1969,  p.  N765)  to  Douville  is  as 
follows:  "The  ancestor  was  a  species  living 
where  its  earliest  representatives  are  found  and 
having  some  of  its  distinctive  characters."  If  one 
uses  this  logical  principle,  then  a  likely  ancestor 
to  Litharca  lithodomus  could  be  Barbatia 
(Cuculla^arca)  reeveana  (Orbigny,  1856)  (plate 
1).  According  to  Keen  (1971),  this  is  a  common 
species  found  throughout  the  present  range  of 
Litharca  lithodomus.  The  ribbing  and  periostra- 
cum  of  Barbatia  (Cucullaearca)  reeveana  resem- 
ble Litharca.  The  size  and  convexity  of  the 
valves  in  the  two  species  are  similar.  There 
would  have  to  be  a  reduction  of  the  byssal  gape 
because  it  is  much  larger  in  Barbatia  (Cucul- 
laearca) reeveana  than  it  is  in  Litharca.  The 
beaks  of  Barbatia  (Cucullaearca)  reeveana  are 
commonly  subcentral  (not  near  the  anterior  end 
as  in  Area),  and  it  would  require  not  a  great 
modification  to  have  the  beaks  thrust  toward 
the  posterior  as  in  Litharca  lithodomus.  Bar- 
batia (CucullcLearca)  reeveana  has  a  habit  of  at- 


taching between  rocks  (Keen,  1971)  so  that  a 
shift  to  a  rock  boring  habit  as  in  Litharca 
lithodomus  would  be  a  likely  change  in  adapta- 
tion. This  is  also  borne  out  by  Yonge's  observa- 
tion (1958)  that  some  rock-boring  species  have 
evolved  from  a  byssate  nestling  species. 

Whether  Litharca  lithodomus  will  be  a  short- 
lived experiment  within  the  Arcidae  or  will  con- 
tinue to  evolve  and  produce  more  rock-boring 
arcids  is  impossible  to  assess.  As  Frizzell 
pointed  out  (1946),  the  lack  of  a  fossil  record  of 
Litharca  lithodomus  may  be  caused  by  the  small 
number  of  rock-boring  habitats  in  the  fossil 
record  and  the  relatively  thin  shells  of  this 
species  inhibiting  preservation.  However,  the 
morphologic  changes  for  the  origin  of  Litharca 
lithodomus  must  have  been  rapid,  and  it  is  quite 
possible  that  the  appearance  of  this  species  was 
less  than  one  million  years  ago. 

Frizzell  (1946)  erected  a  separate  subfamily, 
the  Litharcinae,  for  Litharca  lithodomus,  and 
Yokes  (1980)  has  followed  Frizzell  in  this 
classification.  We  agree  with  Frizzell  and  Yokes 
because  taxonomy  should  show  the  differences 
as  well  as  the  likenesses  between  species,  and 
Litharca  lithodomus  is  so  unique  in  morphology 
and  habit  that  it  merits  being  placed  in  a 
separate  subfamily  within  the  Arcidae. 

Acknowledgment 

We  are  indebted  to  Dr.  Peter  U.  Rodda  of  the 
California  Academy  of  Sciences  for  granting  us 
a  loan  of  specimens  of  Litharca  lithodomus  for 

study. 

LITERATURE  CITED 

Dechaseaux,  C.  1969.  Hippuritoida-Hippuritacea-Origin  and 
extinction,  p.  N765.  In  Treatise  on  Invertebrate  Paleontol- 
ogy. (R.  C.  Moore,  ed.).  Part  N,  Vol.  2,  Mollusca  6, 
Bivalvia.  Univ.  Kansas  Press,  Lawrence. 

Frizzell,  D.  L.  1946.  A  study  of  two  arcid  peleeypod  species 
from  western  South  America.  Jour.  PaLeo.  20:38-51. 

Keen,  A.  M.  1971.  Sea  Shelh  of  Tropical  West  America.  2nd 
ed.  Stanford  Univ.  Press,  California.  1064  p. 

Nicol.  D.  1984.  Changes  in  the  hinge  teeth  of  arcaceans  from 
Devonian  to  Recent.  Tulane  Studies  Geol.  &  Paleo. 
18:70-72. 

Nicol,  D.,  and  D.  S.  Jones.  1984.  Review  of  Postiigata,  a 
Late  Cretaceous  peleeypod.  Tulane  Studies  Geol.  &  Paleo. 
18:67-69. 

Oliver,  P.  (!.  1981.  The  functional  morphology  and  evolution 
of  Recent  Limopsidae  (Bivalvia,  Arcoida).  Malacologia. 
21:61-93. 

Olsson,  A.  A.  1961 .  MoUusks  of  the  Tropical  Eastern  Pacific: 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS    109 


Panamic-Pacific    Pelecypoda.    Paleontological    Research 

Institution.  Ithaca,  New  York.  574  p. 
Rosewater,  J.  1965.  The  family  Tridacnidae  in  the  Indo- 

Pacific.  Indo-Pacific  Mollusca  l:347-.396. 
Stanley,  S.  M.  1972.  Fuctional  morphologj'  and  evolution  of 

byssally    attached    bivalve    mollusks.    Jour.    Paleo. 

46:165-202. 
Thomas,  R.  D.  K.  1976.  Constraints  of  ligamental  growth, 

form  and  function  on  evolution  in  the  Arcidae  (Mollusca: 

Bivalvia).  Paleobiology  2:64-83. 


1978.  Shell  form  and  the  ecological  range  of 

living  and  extinct  Arcoida.  Paleobiolog)'.  4:181-194. 

Yokes,  H.  E.  1980.  Genera  of  the  Bivalvia:  A  Systematic  and 
Bibliographic  Catalogue  (Revised  and  Updated).  Paleon- 
tological  Research  Institution.  Ithaca,  New  York.  307  p. 

Yonge,  C.  M.  1951.  Marine  boring  organisms.  Research 
4:162-167. 

1958.   Observations   on  Petricola   carditoides 

(Conrad).  Proc.  Malaeological  Soc.  London  33:25-31. 


THE  RADULA  OF  ACRORBIS  PETRICOLA 
(PULMONATA:  PLANORBIDAE) 

W.  Lobato  Paraense 

Departamento  de  Malacologia 

Institute  Oswaldo  Cruz 

Caixa  Postal  926 

20000  Rio  de  Janeiro,  RJ-Brasil 

ABSTRACT 
Observation  of  the  radular  teeth  of  Acrorbis  petricola  Odhner,  1937,  and  A. 
odhneri  Scott.  1960  with  the  scanning  electron  microscope  showed  no  difference 
between  the  two  nominal  species,  which  also  are  indistinguishable  in  shell  and 
anatomic  characteristics.  A.  odhneri,  therefore,  is  a  junior  synonym,  of  A. 
petricola.  The  radular  teeth  o/ Acrorbis  are  similar  to  those  o/Gyraulus,  but  taxo- 
nomically  important  differences  in  qualitative  characters  of  the  genital  system 
show  that  the  two  geyiera  are  less  closely  related  than  suggested  by  their  radular 
characteristics. 


Acrorbis  petricola  is  an  atypical  planorbid 
snail  described  by  Odhner  (1937)  from  speci- 
mens from  Nova  Teutonia  (27°16'S,  52°20'W),  a 
village  in  the  Brazilian  state  of  Santa  Catarina, 
sent  to  the  Swedish  Ricksmuseum  by  the 
natural  history  collector,  Herr  Fritz  Plaumann. 
Decisive  characteristics  for  the  erection  of  the 
new  germs  Acrorbis  were  the  Helix-\ike  shape  of 
the  shell,  and  especially  the  squarish  radular 
teeth  with  short  hooks  (see  Fig.  1). 

Pilsbry  (1938)  pointed  to  the  relationship  be- 
tween Acrorbis  and  Drepanotrema  (misprinted 
Drepanostoma)  by  the  similarity  of  the  penial 
complex  and  the  presence  of  a  flagellum  on  the 
penial  sheath,  but  distinguished  the  two  genera 
by  the  teeth  characteristics  as  shown  by  Odhner 
(1937). 

Baker  (1945:123)  commented  on  the  alleged 
differences  in  the  radular  teeth:  "The  radula  [of 


Acrorbis]  appears  very  different  from  that  of 
Drepanotrema  because  of  the  apparent  absence 
of  interstitial  cusps  between  the  larger  cusps. 
These  might  have  been  present  but  overlooked 
because  of  the  small  size  of  the  radula  teeth.  The 
general  shape  of  the  mesocone  in  Odhner's 
figure  4  is  suggestive  of  the  same  cusp  in 
Drepanotrema,  but  the  ectocone  and  entocone 
are  much  shorter."  .  .  .  "Acrorbis.  therefore, 
appears  to  be  another  one  of  the  peculiar  groups 
of  South  American  Planorbidae,  related  to 
Drepanotrema,  but  distinct  by  both  shell  and 
anatomy." 

Examining  specimens  from  Nova  Teutonia, 
Hubendick  (1955)  observed  that  "though  not 
very  detailed,  Odhner's  description  and  figure  of 
the  radula  in  Acrorbis  is  not  incorrect".  His 
drawings  of  a  central,  a  lateral  and  a  marginal 
teeth,  reproduced  in  Fig.  2,  may  be  compared 


no    THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


&&®SFQ 


a^' 


.''^^v^ 


^=^:^ 


^^^^ 


^Tryv^ 


lOjjm 

FIGS.  1-4.  1,  Radular  teeth  of  Acrorbis  petricola  reproduced  from  papers  by  Odhner,  1937;  2,  Hubendick, 
1955;  3,  Scott,  1960,  under  the  name  Acrorbis  odhneri;  4,  Paraense  and  Deslandes,  1959.  (c,  central;  i,  in- 
termediate; 1,  lateral;  m,  marginal.  Scale  bar  refers  to  Fig.  4). 


with  Odhner's  (Fig.  1). 

A  redescription  of  A.  petricola,  based  on 
specimens  collected  at  my  request  by  Herr 
Plaumann  at  the  same  breeding-place  from 
which  the  sample  studied  by  Odhner  had  been 
taken,  was  published  by  Paraense  and 
Deslandes  (1959).  The  teeth  were  represented  as 
shown  in  Fig.  4.  Accessory  cusps  can  be  seen  on 
either  side  of  the  main  cusps  in  the  central  teeth, 
and  also  between  the  two  main  cusps  in  the  fifth 
figured  central  tooth.  Spaces  wider  than  usual 
appear  between  the  bases  of  the  cusps  in  several 
teeth;  such  spaces,  as  later  realized,  were  really 
occupied  by  small  cusps  which  could  not  be 
clearly  discerned  owing  to  their  minuteness 
added  to  light  diffraction  effect.  After  discuss- 
ing the  anatomic  affinities  between  Acrorbis 
and  Drepanotrema,  Paraense  and  Deslandes 
(1959)  concluded  that  the  shape  of  the  shell  was 
the  only  character  that  might  warrant  separa- 
tion of  the  two  genera.  Stating,  however,  that  a 
revision  of  the  genus  Drepanotrema  would  show 
so  wide  interspecific  variation  in  shell  configura- 
tion that  any  generic  diagnosis  based  only  on  the 
shell  characters  of  the  type  species  would  be 
meaningless,  they  treated  A.  petricola.  as  a 
member  of  Drepanotrema.  Subsequently, 
Paraense  (1975)  adopted  the  genus  Acrorbis  on 
account  of  its  Helix-shaped  shell  not  sculptured 


with  longitudinal  rows  of  minute  dots  (puncta- 
tion),  and  of  its  very  long  spiral  radula  project- 
ing far  beyond  the  buccal  mass. 

Under  the  name  Acrorbis  odhneri,  Scott 
(1960)  described  a  snail  from  Argentina  anatom- 
ically identical  with  A.  petricola,  but  differing  in 
shell  characteristics:  larger  size  (largest 
specimen  4.25  mm  in  diameter,  as  against  2  mm 
in  Odhner's  specimens),  rapid  whorl  growth, 
proportionally  wider  aperture,  umbilicus  par- 
tially covered  by  the  basal  reflection  of  the  lip, 
and  absence  of  spiral  lines  on  the  shell  surface. 
Scott  (1960)  states  that  Odhner  so  precisely 
represented  half  a  teeth  row  that  she  could 
recognize  in  it  the  radula  of  her  own  specimens 
(Fig.  3).  (Translated):  "There  are  no  interstitial 
teeth  between  the  cusps  of  the  laterals,  and  the 
marginals  are  markedly  short  showing  no 
oblique  reflection".  Considering  the  lack  of  a 
long  oblique  reflection  in  the  marginals  suffi- 
cient to  separate  Acrorbis  from  the  Planorbidae, 
Scott  (1960)  created  the  family  Acrorbidae  to  in- 
clude A.  petricola  and  A.  odhneri. 

On  17  May  1973  I  collected  23  specimens  of  yl. 
odhneri  at  its  type-locality,  Salto  Encantado 
(27°03'S,  54°50'W),  Departamento  Libertador 
Oeneral  San  Martin,  Province  of  Misiones, 
Argentina.  One  may  easily  reach  Salto  Encan- 
tado by  Road  14  (Ruta  14)  as  far  as  km  212, 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS    111 


where  it  intersects  Ruta  220  (about  14  km  east 
of  Aristobulo  del  Valle,  Departamento 
Cainguas),  and  then  northward  by  the  last- 
mentioned  road  for  about  3  km. 

Salto  Encantado  is  a  waterfall  which  drops 
vertically  and  sprays  the  adjacent  cliff,  favoring 
a  thick  growth  of  moss  where  the  snail  is  found. 
Such  peculiar  habitat  is  similar  to  that  of  A. 
petricola,  which  lives  among  moss  on  rocky 
banks  of  streams  kept  wet  by  trickling  water 
from  the  surrounding  ground  and  above  the 
stream  level. 

The  collected  specimens  were  indistinguish- 
able, in  shell  and  anatomy,  from  A.  petricola 
from  Nova  Teutonia,  as  described  by  Paraense 
and  Deslandes  (1959).  The  largest  shell  was  3.2 
mm  in  diameter.  As  in  the  specimens  from  Nova 
Teutonia  in  our  collection,  the  umbilicus  was  to 
a  variable  extent  covered  by  the  basal  reflection 
of  the  lip  and  the  shell  surface  was  more  or  less 


perceptibly  sculptured  with  thin  spiral  lines. 
Other  differences  mentioned  by  Scott  (1960)  as 
diagnostic  of  A.  petricola  (shell  size,  rate  of 
whorl  increase,  proportion  of  apertural  dia- 
meter) can  be  ascribed  to  intraspecific  or  inter- 
populational  variation. 

As  to  the  radula,  no  significant  difference  was 
observed  between  the  two  forms,  so  that  only 
teeth  of  a  specimen  from  Salto  Encantado  are 
shown  in  Figs.  5-10.  As  observed  with  the  scan- 
ning electron  microscope,  the  central  tooth 
(Figs.  5,  9)  has  two  dagger-like  cusps,  of  which 
the  left  one  is  longer  than  the  right;  there  is  one 
(sometimes  two)  minor  spine-shaped  cusp  high 
on  either  lateral  side  of  the  major  cusps,  and 
another  one  between  the  latter.  The  laterals 
(Fig.  6)  have  three  dagger-like  major  cusps,  of 
which  the  mesocone  is  much  longer,  and  show 
one  smaller  spine-shaped  cusp  high  on  the 
lateral  side  of  the  ectocone  and  entocone  and 


FIGS.  5-lU.  Scanning  electron  micrographs  of  radular  teeth  of  Acrorbis  petricola  from  Salto  Encantado,  Misiones 
province,  Argentina.  5,  central  (arrow),  a  little  slanted  to  the  right;  6,  laterals;  7,  intermediates;  8,  marginals;  9,  central,  a 
little  slanted  to  the  right;  10,  marginals.  Figs.  5-8,  x  29,000;  9,  x  25,000;  10,  x  20,000. 


112    THE  NAUTILUS 


July  30,  1986 


Vol.  100(3) 


between  the  major  cusps.  Small  cusps  between 
and  outside  the  three  major  cusps  are  added 
from  the  intermediates  (Fig.  3)  to  the  extreme 
marginals  (Figs.  4,  6),  while  the  ectocone,  the 
entocone  and  particularly  the  mesocone  remain 
easily  distinguishable.  Both  major  and  minor 
cusps  grow  smaller  toward  the  edges  of  the 
radular  ribbon.  The  marginals  have  short  wide 
reflections  high  up  on  the  base  of  attachment. 
The  extreme  marginals  usually  show  13-14 
cusps,  including  the  three  main  ones. 

The  radular  teeth  of  Acrorbis  are  similar  to 
those  of  Gyraulus,  recently  studied  by  Meier- 
Brook  (1983)  and  Burch  and  Jeong  (1984),  but 
the  absence  in  Acrorbis  of  taxonomically  impor- 
tant characters  such  as  a  penial  stylet  and  a 
separate  prostatic  duct,  and  the  presence  of 
flagella  on  the  penial  sheath,  show  that  the  two 
genera  are  less  closely  related  than  suggested 
by  their  radular  characteristics. 

The  observations  described  above  point  to  the 
identity  of  A.  petricola  and  A.  odhneri,  lending 
no  support  to  the  removal  of  the  genus  from  the 
family  Planorbidae. 

Acknowledgments 

The  author  is  grateful  to  Petrobras-CENPES 


(Centro  de  Pesquisas  e  Desenvolvimento)  for 
use  of  the  facilities  of  its  SEM  Laboratory,  to 
Carlos  R.  Cordeiro  and  Flavio  J.  Versiani  for 
technical  assistance,  and  to  Prof.  Walter  Berto- 
lazzo,  Universidade  Federal  do  Rio  de  Janeiro, 
who  introduced  the  author  to  the  staff  of  the 
SEM  Laboratory. 


LITERATURE  CITED 

Baker,  F.  C.  1945.  The  molluscan  family  Planorbidae.  Univ. 

Illinois  Press,  Urbana.  xxxvi  +  530  p. 
Burch,  J.  B.  and  K.  H.  Jeong.  1984.  The  radular  teeth  of 

selected  Planorbidae.  Malacol.  Rev.  17:67-84. 
Hubendick,  B.  1955.  Phylogeny  in  the  Planorbidae.  Trans. 

Zool.  Soc.  London  28:453-542. 
Meier-Brook,    C.    1983.   Taxonomic   studies   on   Gyraulus 

(Gastropoda:  Planorbidae).  Malacologia  24:1-113. 
Odhner,  N.  H.  1937.  Acrorbis  petricola  n.  gen.  n.  sp.,  eine 

merkliche  Siisswasserschnecke  aus  Brasilien.  Ark.  Zool. 

29B:l-8. 
Paraense,  W.   L.   1975.  Estado  atual  da  sistematica  dos 

planorbideos  brasileiros.  Arq.  Mus.  Nac.  Rio  de  Janeiro, 

55:105-128. 
Paraense,  W.  L.  and  N.  Deslandes.  1959.  The  Brazilian 

species  oi  Drepanotrema.  VII.  D.  petricola  (Odhner,  1937). 

Rev.  Brasil  Biol.  19:319-329. 
Pilsbry,    H.    A.    1938.   Acrorbis   petricola.    Review.    The 

Nautilus  51:107. 
Scott,  M.  I.  H.  1960.  Nueva  familia  de  pulmonado  basoma- 

toforo  (Mollusca).  Neotropica  6:65-69. 


NATICA  (GLYPHEPITHEMA)  TEDBAYERI,  A  REPLACEMENT  NAME 

Harald  A.  Rehder 

National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


In  a  recent  issue  of  The  Nautilus  (Rehder, 
1986:  Nautilus,  100(1):38)  I  proposed  the  name 
Natica  (Glyphepithema)  bayeri  for  Glyphe- 
pithema  Jloridana  Rehder,  1943,  not  Natica 
Jloridana  Dall,  1892. 

Now  Alan  R.  Kabat  at  the  Museum  of  Com- 
parative Zoology  has  kindly  called  my  attention 
to  the  fact  that  the  taxon  Natica  bayeri  is  pre- 


occupied by  Natica  (Naticina)  bayeri  Koper- 
berg,  1931  (Jaarboek  von  het  Mijnwezen  in 
Nederlandsch  Oost-Indie,  59:  139,  pi.  3,  fig.  48) 
from  the  Pliocene  of  Timor,  Indonesia. 

I  propose  now  to  name  the  species  found,  from 
southeastern  Florida  to  Brasil,  Natica 
(Glyphepithema)  tedbayeri  Rehder. 


Vol.  100(3) 


July  30,  1986 


THE  NAUTILUS    113 


BOOK  REVIEWS 
Seashells  of  Western  Australia  by  Fred  E.  Wells 
and  Clayton  W.  Bryce.  1986.  Western  Austra- 
lian Museum,  Perth.  207  pp.,  74  colored  pis. 
Paperback.  $17.95  Australian  (approx.  U.S. 
$12.50). 

This  is  an  excellent,  well-illustrated  guide  to 
the  more  common  and  readily  found  species  of 
marine  mollusks  of  Western  Australia.  The 
western  third  of  Australia  is  particularly  rich  in 
molluscan  species  because  of  the  meeting  of  the 
tropical  Indo-Pacific  province  in  the  north  with 
the  colder  temperate  waters  of  South  Australia 
and  Tasmania.  Of  the  approximate  2000  species 
from  this  region,  this  accurate  guide  beautifully 
illustrates  671  species,  including  many  photo- 
graphs of  living  specimens  of  opisthobranchs 
and  cephalopods.  Each  family  is  defined  and, 
when  available,  useful  bibliographic  references 
are  included.  Scientific  name,  author  and  date, 
size,  geographical  distribution,  relative  abun- 
dance within  Western  Australia,  and  an  ex- 
cellent photograph  are  given  for  each  of  these 
common  and  conspicuous  species.  A  short  in- 
troduction on  protecting  mollusks,  cleaning  and 
maintaining  a  collection  of  shells,  together  with 
a  glossary  and  index  round  out  this  fine  little 
book.-/?.  T.  Abbott. 

The  Littorinid  Molluscs  of  Mangrove  Forests  in 
the  Indo-Pacific  Region,  by  David  G.  Reid.  24 
April  1986.  228  pp.,  98  figs.,  1  color  plate. 
Publ.  no.  978,  British  Museum  (Natural 
History).  Cloth,  $35.00  (Approximately  U.S. 
$52.00). 

This  is  one  of  the  most  complete  and  com- 
petently executed  monographs  of  a  well-known 
group  of  common  mangrove-dwelling  gastro- 
pods. The  taxonomy  of  this  otherwise  perplex- 
ing genus,  Littoraria.  has  been  well  worked  out. 
Despite  the  recognition  of  27  Indo-Pacific 
species  and  subspecies  in  a  group  heretofore 
usually  treated  as  three  or  four  good  species,  the 
author  has  proposed  only  two  Indo-Pacific  new 
species,  one  new  subspecies  and  one  new  sub- 
genus, (Paliistorina).  His  detailed  presentation 
of  anatomical,  egg-capsule,  radular  and  distribu- 
tional characteristics  will  probably  lead  to  a 
universal  acceptance  of  his  views.  Very  for- 
tunately a  large  number  of  earlier  names  solely 
based  on  shell  characters  were  available  for 


these  new  biologically  recognized  species.  Ex- 
cellent drawings  of  penes,  pallial  oviducts, 
sperm  nurse  cells,  and  distributional  maps  ac- 
company each  of  the  27  Indo-Pacific  taxa. 
Worldwide,  37  taxa  are  recognized  in  five  sub- 
genera of  the  genus  Littoraria  Griffith  and 
Pidgeon,  1834.  The  first  70  pages  of  this  work 
have  an  excellent  discussion  of  morphological 
characters,  reproduction,  habitats  and  biogeo- 
graphy.-/?.  T.  Abbott. 

It's  Easy  to  Say  Crepidula!  by  Jean  M.  Cate  and 
Selma  Raskin.  1986.  155  pp.  Pretty  Penny 
Press.  P.O.  Box  3890,  Santa  Monica,  CA 
90403.  Paperback,  $19.95,  plus  $1.50  postage. 

This  is  a  useful  phoenetic  guide  to  the  pronun- 
ciation of  the  scientific  names  of  about  3,000 
common  molluscan  taxa.  The  authors  usually 
give  two  pronunciations,  one  evidently  used  in 
American  Latin  classrooms,  the  other  used  cus- 
tomarily among  most  American  malacologists. 
Readers  may  take  their  choice.  However,  the 
customary  pronunciation  of  the  -ae  ending,  used 
in  species  named  for  a  woman,  is  not  given.  The 
name  myrakeeyiae,  for  instance,  is  usually  pro- 
nounced: MY'  ruh  KEEN'  ee,  but  the  authors 
give  only  MY'  ruh  KEEN'  ay  and  MY'  ru  KEEN' 
eye.  They  are  correct  in  suggesting  that  names 
for  males,  ending  in  -i,  may  be  pronounced 
"eye",  as  in  laurenti:  law  RENT'  eye.  There  is 
also  an  eleven-page  glossary  of  malacological 
terms.-/?.  T.  Abbott. 

Seashell  Treasures  of  the  Caribbean  by  Lesley 
Sutty.  Edited  by  R.  Tucker  Abbott.  E.  P. 
Dutton,  New  York.  1986.  128  pp.,  139  color 
plates.  $19.95.  (with  autographed,  numbered 
bookplate  $21.95  from  American  Malacolo- 
gists, P.O.  Box  1192,  Burlington,  MA  01803). 

The  author,  an  accomplished  naturalist,  diver 
and  photographer,  has  chosen  100  species  of 
Caribbean  mollusks  "to  illustrate  and  comment 
upon  so  that  the  reader  may  share  my  enthu- 
siasm and  joys  in  discovering  the  shells  of  these 
beautiful  tropical  islands."  She  has  attempted  to 
inspire  others  to  study  mollusks  by  relating  her 
experiences  in  seeking  these  elusive  and  fascin- 
ating animals  in  the  Lesser  Antilles.  Her  lively 
and  engaging  narrative,  coupled  with  her  stun- 
ning photographs,  are  certain  to  provide  any 
shell   lover   with   many   hours   of  pleasurable 


114    THE  NAUTILUS 


July  30,  1986 


Vol.  100  (3) 


reading,  and  should  stimulate  interest  in  seek- 
ing and  observing  marine  shells. 

A  preface  by  the  editor  and  the  author's  in- 
troduction begin  this  volume,  followed  by 
chapters  entitled,  "What  is  a  Rare  Shell?", 
"Strange  and  Beautiful  Shapes",  "Treasures  of 
Sea  and  Reef,  and  "Science  in  Shells."  Rare 
shells  discussed  and  illustrated  include  Cypraea 
surinamensis,  Pterynotus  phyllopterus  and 
Conies  granulatus,  to  name  just  a  few.  Unusual 
mollusks,  such  as  Umbraculum  umbraculum, 
Glossodoris  clenchi  and  Xenophora  conchylio- 
phorci  become  familiar  through  the  author's 
animated  writing  style.  The  bulk  of  the  species 
are  treated  in  the  "Treasures"  chapter  where  we 
meet  the  rediscovery  of  Hexaplex  straussi  and 
such  desirable  species  as  Lyria  archeri,  Conus 
cedonulli  and  Chlamys  multisquamata.  An  ab- 
breviated history  of  Caribbean  malacology, 
followed  by  a  brief  bibliography,  index  and 
glossary  of  terms,  completes  the  text. 

There  is  a  great  deal  of  merit  in  this  beautiful- 
ly illustrated  book,  and  it  is  important  again  to 
stress  that  the  author  has  fully  achieved  her  aim 
of  sharing  with  others  her  collecting  activities. 
This  book  is  not  intended  to  be  an  identification 
guide.  Despite  meticulous  editing,  last  minute 
publisher's  errors  do  creep  in,  such  as  the 
transposition  of  the  captions  for  plates  22  and 
23,  and  the  indiscriminate  use  of  parentheses 


around  every  author  and  date.  Nonetheless,  it  is 
hoped  that  this  well-produced  and  lavishly  il- 
lustrated work  will  be  enjoyed  and  appreciated 
by  nature  lovers  and  conchologists,  and  will 
serve  to  introduce  living  mollusks  to  many  new 
enthusiasts -Walter  E.  Sage. 

North  Atlantic  Nudibranchs  (MoUusca)  Seen  By 
Henning  Lemche  by  Hanne  Just  and  Malcolm 
Edmunds.   1985.    170  pp.,   69  pis.   in  color. 
Paperback.  Supplement  2  of  Ophelia,  Inter- 
national Journal  of  Marine  Biology,  Denmark. 
500  kroner  (approx.  U.S.  $60.00) 
The  excellent  69  color  plates  depicting  the  late 
Dr.  Henning  Lemche's  personal  paintings  of  76 
of  the  known  217  species  of  opisthobranchs  of 
the  North  Atlantic  will  prove  useful  to  students 
of  nudibranchs.  The  descriptions  of  the  species 
and  the  information  on  feeding  and  reproduc- 
tion, together  with  up-dated  taxonomic  notes  by 
the  author-editors  add  considerable  usefulness 
to  the  book.  A  geographical  checklist  of  the  217 
species  by  Elizabeth  Platts  is  very  helpful. 

Most  of  Lemche's  illustrated  species  are  in  the 
genera  Doto,  Onchidoris,  Flabellina.  Eubran- 
chus,  Cuthona  and  Aeolidiella.  26  of  the  76 
species  are  given  no  trivial  names  and  await  fur- 
ther taxonomic  consideration.  Perhaps  someday 
this  unfinished  book  will  be  carried  to  conclusion 
by  one  or  both  of  the  authors. -i?.  T.  Abbott. 


INFORMATION  FOR  SUBSCRIBERS 


The  annual  sviscription  rate  for  The  Nautilus 
is  $15.00  for  individuals  (foreign  $18.00)  and 
$20.00  for  institutions  (domestic  or  foreign). 
Subscriptions  may  begin  in  January.  Send  check 
or  money  order  made  out  to  "American  Mala- 
cologists"  to  the  Business  Manager,  P.O.  Box 
2255,  Melbourne,  Florida  32902-2255,  U.S.A. 


Back  issues  from  volume  72  to  date  are  ob- 
tainable from  the  Business  Manager.  Volumes  1 
through  71  (if  available)  may  be  obtained  in 
reprint  or  original  form  from  Kraus  Reprint 
Co.,  Route  100,  Millwood,  New  York  10546. 

Advertising  rates  may  be  obtained  from  the 
Business  Manager  or  Editor. 


CONTRIBUTORS 


Mamcscripts:  Authors  are  requested  to  follow 
the  recommendations  of  the  Style  Manual  for 
Biological  Journals,  which  may  be  purchased 
from  the  American  Institute  of  Biological  Sci- 
ences, 1401  Wilson  Boulevard,  Arlington,  Va. 
22209.  Manuscripts  should  be  typewritten  and 
doublespaced;  original  and  one  copy  are  re- 
quired, to  facilitate  reviews.  Tables,  numbered 
in  arable,  should  be  on  separate  pages,  with  the 
title  at  the  top.  Legends  to  photographs  should 
be  typed  on  separate  sheets.  Explanatory  terms 
and  symbols  within  a  drawing  should  be  neatly 
printed,  or  they  may  be  pencilled  in  on  a  translu- 


cent overlay,  so  that  the  printer  may  set  them  in 
8  pt.  type.  There  is  a  charge  of  50  cents  per 
word  for  this  extra  service.  All  authors  or  their 
institutions  will  be  charged  50  cents  per  line  of 
tabular  material  and  taxonomic  keys.  The  pub- 
lishers reserve  the  right,  seldom  exercised,  to 
charge  $45  per  printed  page. 

An  abstract  should  accompany  each  paper. 

Reprints  are  available  at  cost  to  authors. 
When  proof  is  returned  to  authors,  information 
about  ordering  reprints  will  be  given.  They 
are  obtained  from  Economy  Printing  Co.,  Inc., 
R.D.  3,  Box  169,  Easton,  Maryland  21601-9430. 


MOLLUSK  VOUCHER  SPECIMENS 


It  is  becoming  increasingly  important  for 
future  research  purposes  that  an  identified  sam- 
pling of  species  mentioned  in  publications  be 
deposited  in  a  permanent,  accessible  museum 
specializing  in  mollusks.  This  is  particularly  true 
of  mollusks  used  in  physiological,  medical, 
parasitological,  ecological,  and  experimental 
projects. 


Several  museums  of  natural  history  have  ex- 
tensive modern  facilities  and  equipment  for  the 
housing  and  curating  of  voucher  specimens. 
Material  should  be  accompanied  by  the  identifi- 
cation, locality  data  and  its  bibliographic 
reference.  There  is  no  charge  for  this  perma- 
nent curating  service,  and  catalog  numbers,  if 
desired,  will  be  sent  to  authors  prior  to  publica- 
tion. 


WANTED  -  OLD  SHELL  BOOKS 

Will  pay  good  prices  for  libraries,  second-  Phone  (1-305-725-2260)  or  write:  R.  Tucker 

hand  books  and  reprints  on  mollusks,  shells  Abbott,  American  Malacologists,  Inc.,  P.O. 

and    conchology.    Back    numbers    of    The  Box  2255,  Melbourne,  FL  32902.  Free  ap- 

Nautilu^,  vols.  40-71  wanted,  $1.50  each.  praisals. 


americanmalacologists,  inc. 

PUBLISHERS  OF  DISTINCTIVE  BOOKS  ON  MOLLUSKS 

THE  NAUTILUS  (Quarterly) 

MONOGRAPHS  OF  MARINE  MOLLUSCA 

STANDARD  CATALOG  OF  SHELLS 

INDEXES  TO  THE  NAUTILUS 

{Geographical,  vols  1-90;  Scientific  Names,  vols  61-90) 

REGISTER  OF  AMERICAN  MALACOLOGISTS 


OCTOBER  31,  1986 


THE 


NAUTILUS 


ISSN  0028-1344 


Vol.  100 


No.  4 


A  quarterly 

devoted  to 

malacology  and 

the  interests  of 

conchologists 


UBRAnr 

NOV    4  1986 

-- ^SlHoie,  Mass. 


Founded  1889  by  Henry  A.  Pilsbry.  Continued  by  H.  Burrington  Baker. 
Editor-in-Chief:  R.  Tucker  Abbott 


EDITORIAL  COMMITTEE 
CONSULTING  EDITORS 


Dr.  William  K.  Emerson 
Department  of  Living  Invertebrates 
The  American  Museum  of  Natural  History 
New  York,  NY  10024 

Mr.  Samuel  L.  B.  Fuller 
1053  Mapleton  Avenue 
Suffield,  CT  06078 

Dr.  M.  G.  Harasewych 
363  Crescendo  Way 
Silver  Spring,  MD  20901 

Mr.  Richard  L  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Los  Angeles  County  Museum  of  Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 


Dr.  Arthur  S.  Merrill 
c/o  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Donald  R.  Moore 
Division  of  Marine  Geology 
School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 

Dr.  G.  Alan  Solem 
Department  of  Invertebrates 
Field  Museum  of  Natural  History 
Chicago,  IL  60605 

Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Cambridge,  MA  02138 

Dr.  Gilbert  L.  Voss 
Division  of  Biology 

School  of  Marine  and  Atmospheric  Science 
10  Rickenbacker  Causeway 
Miami,  FL  33149 


EDITOR-IN-CHIEF 

Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
Box  2255,  Melbourne,  FL  32902-2255 

Mrs.  Cecelia  W.  Abbott 
Business  and  Subscription  Manager 
P.O.  Box  2255 
Melbourne,  FL  32902-2255 


Second  Class  Postage  paid  at  Melbourne,  Florida 
and  other  post  offices 


The  Nautilus  (USPS  374-980) 
ISSN  0028-1344 

A  quarterly  magazine  devoted  to  malacology. 
Copyright  ^^1986  by  American  Malacologists,  Inc. 

OFFICE  OF  PUBLICATION 

American  Malacologists,  Inc.  (United  Parcel  Address: 

2208  South  Colonial  Drive,  Melbourne,  FL  32901) 

Mail:  Box  2255,  Melbourne,  FL  32902-2255 

POSTMASTER:  Send  address  changes  to  above. 

Subscription  Price:  $15.00  (see  inside  back  cover) 

$17.00  (foreign);  institutions  $20.00 


Marine  BioioQical  bboralorv 
LIBRARY 

I        NOV    4  19B6 

Woods  Hole,  Mass. 


THE 

NAUTILUS 

Volume  100,  number  4  -  October  31,  1986 

ISSN  0028-1344 


CONTENTS      " 

Louise  Russert-Kraemer  and  Elmer  G.  Berry 

"Things  Truly  Excellent.  .  .":  Henry  Van  Der  Schalie,  1907-1986 116 

R.  Tucker  Abbott 

Cauthdt'us  multangulus  New  Subspecies  Grandanus  from 

Northwest  Florida  (Buccinidae) 120 

Jon-Arne  Sneli  and  Oystein  Stokland 

On  the  Taxonomical  Status  of  Tritonium  viridulum  Fabricius,  1780 

(Gastropoda:   Cancellariidae) 121 

Douglas  G.  Smith 

Pisidium  henslowanum  (Sheppard)  in  the  Connecticut  River, 

Massachusetts  (Bivalvia:  Pisidiidae) 124 

C.  M.  Hawkins  and  R.  B.  Angus 

Preliminary  Observations  of  Predation  on  Ocean  Quahaugs,  Arctictt  islandica, 

by  Atlantic  Wolffish,  Annrhichas  lupu^; 126 

Jose  D.  Gomez.  Mercedes  Vargas  and  Emile  A.  Malek 

Freshwater  Mollusks  of  the  Dominican  Republic 130 

Paul  W.  Parmalee  and  Walter  E.  Klippel 

A  Prehistoric  Aboriginal  Freshwater  Mussel  Assemblage  from 

the  Duck  River  in  Middle  Tennessee 134 

David  Nicol 

The  Fate  of  Pelecypod  Families,  Subfamilies,  and  Tribes  During 

and  After  the  Cretaceous  Period 140 

Emile  A.  Malek 

Freshwater  and  Terrestrial  Snails  of  Saint  Lucia,  West  Indies 143 

William  K.  Emerson  and  Walter  E.  Sage  III 

On  the  Rediscovery  of  Teniniiiehin  minihilin  (Clench  and  Aguayo,  1941), 

and  Its  Relationship  to  Other  Calliotectine  Volutes 147 

To  Joe  Rosewater  — from  R.  Tucker  Abbott 

Reminiscences 152 


All  future  subscriptions  and  manuscripts  should  be  sent  to  Tin 
Nautilus.  P.O.  Box  3430,  Silver  Spring,  MD  20901,  U.S.A. 


116    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


"THINGS  TRULY  EXCELLENT.  .  .":  HENRY  VAN  DER  SCHALIE, 

1907-1986 

Louise  Russert-Kraemer  and  Elmer  G.  Berrv 


Henry  van  der  Schalie  (1907-1986) 

Henry  van  der  Schalie,  extraordinary  scholar, 
teacher  and  friend  to  so  many  of  us,  died  Tues- 
day afternoon,  15  April,  1986,  in  his  home  near 
the  Raisin  River  in  Manchester,  Michigan,  with 
his  beloved  wife  and  colleague,  Annette 
Rudolphi  van  der  Schalie,  at  his  side.  Born  in 
1907  in  Amsterdam,  the  Netherlands,  Professor 
van  der  Schalie  completed  high  school  in  Pater- 
son,  New  Jersey,  and  then  earned  an  A.B. 
degree  from  Calvin  College  in  Minnesota  in 
1929,  an  M.S.  degree  from  the  University  of 
Michigan  in  1931,  and  a  Ph.D.  from  the  Univer- 
sity of  Michigan  in  1934.  At  the  University  of 
Michigan,  he  served  from  1929  to  1934  as 
Museum  Assistant,  from  1934  to  1944  as  Assis- 


tant Curator  of  Mollusks,  and  as  Curator  of 
Mollusks  from  1944  to  1977.  He  also  held  an  ap- 
pointment as  Instructor  of  Zoology  at  the 
University  of  Michigan  from  1934  to  1937,  and 
served  as  Exchange  Professor  at  the  University 
of  Puerto  Rico  from  1940  to  1944.  Returning  to 
the  University  of  Michigan,  he  became  Assistant 
Professor  of  Zoology  in  1944,  Associate  Pro- 
fessor in  1950,  and  Professor  of  Zoology  in  1957, 
a  position  he  held  until  his  retirement  in  1977. 

Highly  respected  as  an  outstanding  interna- 
tional authority  on  mollusks,  Professor  van  der 
Schalie  served  on  numerous  committees  and  as 
consultant  to  government  agencies  in  Egypt, 
Sudan  and  Japan.  He  served  as  a  consultant  to 
the  World  Health  Organization,  as  well.  His 
work  for  these  organizations  centered  on  the 
mollusk  hosts  of  human  pathogens,  such  as 
Schistosoma.  A  well-known  authority  not  only 
on  disease-bearing  snails  around  the  world,  but 
on  land  and  freshwater  mollusks  of  Puerto  Rico, 
on  the  value  of  mussel  distribution  in  tracing 
stream  confluence,  on  the  biology  of  land  and 
freshwater  mollusks,  on  medical  malacology,  on 
paleoecology,  and  other  subjects  as  well.  Pro- 
fessor van  der  Schalie  published  well  over  110 
manuscripts  on  his  research.  More  complete 
details  on  his  professional  activities  can  be 
found  in  American  Malacologists  and  its  sequel, 
the  Register  of  American  Malacologists  (1986). 

In  the  Bulletin  of  the  American  Malacological 
Union  for  1980,  Professor  van  der  Schalie  pro- 
vided an  illuminating  review  of  "Fifty  years  of 
Malacology  at  the  University  of  Michigan  (1929- 
1975)."  Herein  he  summarized  the  outstanding 
features  of  the  "Goodrich  Period  (1929-1944)", 
the  "van  der  Schalie  Period  (1944-1977)"  and  the 
"Highlights  of  the  Mollusk  Collections"  during 
those  years  at  the  Mollusk  Division  of  the 
Museum  of  Zoology  at  the  University  of 
Michigan. 

At  a  festschrift  organized  by  his  students,  for 
Professor  van  der  Schalie  in  Ann  Arbor  in  April, 
1978,  on  the  occasion  of  his  retirement  in  1977, 
many   tributes   were   presented    to   "Van"   by 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS     117 


dozens  of  his  students,  colleagues  and  friends. 
Of  particular  poignance  was  the  careful  com- 
ment offered  by  one  of  Van's  oldest  friends  and 
colleagues.  Professor  Elmer  G.  Berry.  Several 
paragraphs  from  Dr.  Berry's  unpublished 
reminiscence  on  that  occasion,  follow: 

"Henry  van  der  Schalie  was  born  in  Amster- 
dam, January  8,  1907.  His  father  died  when 
Henry  was  but  an  infant  and  his  mother,  in 
1909,  emigrated  from  Holland  to  Paterson,  New 
Jersey,  with  Henry  and  his  two  older  brothers. 
Circumstances  necessitated  Mrs.  van  der 
Schalie  to  find  employment  to  provide  for 
herself  and  the  three  boys,  John,  Herman  and 
Henry.  She  was  very  gifted  in  knitting, 
crocheting  and  sewing  in  general  and  found 
employment  in  a  cotton  factory  where,  among 
other  items,  skirts  were  manufactured.  Most 
likely,  all  three  boys  worked  in  this  plant  after 
school  and  during  the  summer  months.  Henry 
was  assigned  to  the  packing  department,  par- 
ticularly the  wrapping  and  tying  of  cartons  of 
shirts  for  shipment.  Anyone  who  has  observed 
Henry  wrapping  and  tying  a  carton  can  vouch 
for  his  adroitness  from  the  experience  gained 
when  he  was  a  mere  lad. 

"Somewhere  around  1925  Henry  left  Paterson 
for  Grand  Rapids,  Michigan,  where  he  enrolled 
as  a  student  in  Calvin  College.  Although  he 
majored  in  Zoology  he  also  completed  the  pre- 
medical  requirements  because  of  his  indecision 
regarding  his  future  profession.  The  head  of  the 
Department  at  Calvin  College  was  Dr.  van 
Heisman,  a  recognized  parasitologist.  Henry 
had  been  influenced  by  (Dr.  Heisman)  and  began 
to  concentrate  in  the  field  of  parasitology.  His 
splendid  scholastic  record  during  his  under- 
graduate years  merited  commendable  recom- 
mendations from  his  professors.  Although  four 
universities  offered  him  assistantships,  he  chose 
the  University  of  Michigan  and  began  his  work 
under  Dr.  George  R.  LaRue,  an  outstanding 
parasitologist  and  Chairman  of  the  Department 
of  Zoology. 

(After  coming  to  Ann  Arbor,  Henry  accepted 
an  assistantship  in  the  Division  of  Mollusks  in 
the  new  Museum  of  Zoology.  With  some  reluc- 
tance, Dr.  Berry  reports.  Dr.  La  Rue  allowed 
Henry  to  transfer.) 

"Calvin  Goodrich  was  the  Curator  of  Mollusks 
at  that  time.  He  had  formerly  been  the  editor  of 


a  newspaper  in  Ohio,  but  had  taken  an  interest 
in  collecting  shells  as  a  hobby.  He  was  ac- 
quainted with  Dr.  Bryant  Walker,  a  highly  suc- 
cessful practicing  lawyer  in  Detroit.  Walker  had 
made  an  outstanding  reputation  as  a  mala- 
cologist  and  had  been  appointed  Honorary 
Curator  of  Mollusks  at  the  Museum  of  Zoology. 
Goodrich's  association  with  Dr.  Walker  brought 
him  in  contact  with  A.  G.  Ruthven,  Director  of 
the  University  Museums  (later  President  of  the 
University  of  Michigan)  and  with  F.  M.  Gaige, 
Director  of  the  Museum  of  Zoology.  It  is 
reported  that  Goodrich  asked  Ruthven  if  he 
could  have  a  small  corner  in  the  new  Museums 
to  "play  with  his  shell  collection."  He  was  given  a 
room,  but  the  relationship  between  him  and  the 
fine  Curator  of  Mollusks  was  not  the  best,  which 
subsequently  resulted  in  the  resignation  of  Miss 
Mina  Winslow.  With  this  termination,  Mr. 
Goodrich  was  appointed  Curator. 

"Goodrich  was  a  highly  sensitive  individual 
and  unquestionably  realized  that  being  ap- 
pointed Curator  of  Mollusks  at  the  University  of 
Michigan  was  a  unique  position  with  a  (status) 
equal  to  that  of  Harvard  or  the  Philadelphia 
Academy  of  Natural  Sciences.  He  was  not 
academically  trained  as  a  scientist  (and  that) 
caused  him  to  develop  an  inferiority  complex. 
He  had  a  flair  for  writing,  especially  historical  or 
narrative  experiences  such  as  a  collecting  trip, 
and  was  a  much  better  editor  of  his  newspaper 
than  a  curator  of  mollusks. 

Elmer  Berry  continues:  "Mr.  Goodrich  replied 
to  my  application  for  an  assistantship  (at  the 
Museum  in  1933)  with  a  frank  statement  that 
the  Museum's  budget  had  already  been  trim- 
med. He  advised  me  that  the  only  job  likely  to  be 
open  would  consist  of  feeding  animals  and  clean- 
ing their  quarters.  With  the  severe  cut  of  ap- 
propriations, even  this  job  might  be  eliminated 
because  of  lack  of  funds.  (In)  his  letter  written  to 
me  on  February  2,  1933,  he  wrote,  'We  have 
been  rather  fortunate  in  the  students  who  have 
worked  in  this  division -H.B.  Baker,  whose 
work  you  are  familiar  with;  W.  J.  Clench  who  is 
now  clearing  up  his  long  studies  of  the  Physidae; 
Henry  van  der  Schalie  promising  to  be  an 
authority  on  the  Unionidae;  and  Alan  Archer, 
who  already  knows  land  shells  very  well  and  has 
undertaken  to  straighten  out  the  Viviparidae  of 
North  America.  It  would  be  a  pleasure  to  add 


118    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


your  name  to  this  small,  but  excellent,  company, 
and  my  hope  is  that  it  can  be  brought  about.' 

"I  shall  always  remember  my  introduction  to 
Ann  Arbor  and  the  Museum  of  Zoology.  It  was 
terribly  hot  and  humid,  (July,  1933)  but  I  was 
impressed  with  the  beautiful  rotunda  of  the 
Museum's  building  and  the  graceful  marble 
stairs  leading  to  the  second  floor.  That  morning 
I  met  Henry  van  der  Schalie.  He  was  very 
friendly  and  about  the  first  thing  he  asked  was 
where  I  was  staying.  When  I  told  him,  the 
Allenel  Hotel,  he  replied,  'You  can't  afford  to 
stay  there  very  long!'  Henry  had  been  raised  to 
be  thrifty  as  well  as  energetic.  Each  month  a 
portion  of  his  small  salary  went  to  his  mother 
and  the  remainder  paid  for  clothing,  tuition, 
books,  room  and  board.  He  learned  quickly  that 
my  salary  of  $60  per  month  would  necessitate 
stringent  budgeting  because  the  cost  of  living  in 
Ann  Arbor,  even  in  1933,  was  very  high.  He  in- 
vited me  to  have  lunch  with  him,  providing  I 
could  get  along  on  a  peanut  butter  and  jelly 
sandwich,  which  I  was  happy  to  accept.  In  order 
to  cut  expenses  Henry  and  three  other  zoolo- 
gists were  sharing  a  two-room  apartment  just  a 
half-block  east  of  the  Museum  Building  on  North 
University  Street.  I  don't  know  what  persuasive 
powers  Henry  exerted  on  his  three  roommates, 
but  by  two  o'clock  on  my  first  day  in  Ann  Arbor, 
I  was  invited  to  become  the  fifth  member  of  this 
fraternity.  Henry's  Dutch  ancestry  and  home- 
training  instilled  in  him  a  delight  in  getting 
things  clean.  Each  of  us  had  certain  duties  to 
perform  in  the  apartment  and  Henry  was 
assigned  to  cleaning  the  kitchen  and,  on  alter- 
nate Saturdays,  the  bathroom.  On  reciprocal 
Saturdays  the  bathroom  was  to  be  cleaned  by 
Nap  Curtis  who  lived  in  the  west  wing  of  the  2nd 
floor.  When  Henry  finished  his  assignment,  the 
bathroom  sparkled  with  cleanliness.  Nap,  in  con- 
trast, considered  it  unnecessary  to  spend  much 
time  in  cleaning.  As  soon  as  Henry  discovered 
(Nap's)  perfunctory  job,  a  furor  developed  and 
eventually  provoked  a  breach  of  friendship  be- 
tween the  two. 

"Henry  received  his  I'h.U.  in  1934  and  at  the 
beginning  of  the  new  fiscal  year  he  was  ap- 
[)ointed  Assistant  Curator  of  Mollusks.  The 
American  Malacological  Union  meetings  were 
scheduled  to  be  held  at  Stanford  University  dur- 
ing the  summer.  Coodrich,  who  was  one  of  the 


founders  of  the  Union,  gave  his  endorsement 
(for  Henry  to  attend,  and  present  a  paper  on  his 
dissertation).  I  was  invited  to  drive  west  with 
him  in  his  Model  A  Ford.  It  was  Henry's  first 
trip  west  so  we  made  plans  to  see  as  much  as  we 
could  in  the  limited  time.  We  included  Yellow- 
stone, the  Tetons,  Great  Salt  Lake  and  planned 
to  do  some  collecting  in  the  western  canyons, 
Pyramid  Lake -a  relic  lake  in  Nevada,  Lake 
Tahoe,  as  well  as  some  side  trips  in  California. 
We  had  a  delightful  trip  and  met  many  mala- 
cologists  for  the  first  time,  the  Junius  Hender- 
sons, the  Oldroyds,  and  saw  a  few  old  friends, 
Stillman  Berry,  Allyn  Smith,  Leo  Hertlein  and 
others.  Best  of  all  Henry  and  I  got  to  know  each 
other  very  well  and  found  each  other's  company 
very  compatible. 

"Late  in  1935  Henry  became  an  active  mem- 
ber of  Phi  Sigma.  He  enjoyed  the  meetings  and 
found  the  members  very  congenial.  Particularly, 
he  was  attracted  to  a  botanist  by  the  name  of 
Annette  Rudolphi.  Both  of  them  enjoyed  ice 
skating  and  taking  walks  through  the  arbore- 
tum. By  the  spring  of  1936  it  was  obvious  to  me 
that  this  attraction  had  developed  into  a  serious 
romance.  Walking  to  the  Museum  and  back 
again,  or  while  preparing  dinner  or  washing 
dishes,  the  major  conversation  was  about  his 
girl  friend.  By  midsummer  Henry  informed  me 
that  I  had  better  find  another  place  to  live 
because  they  were  going  to  be  married  and 
would  occupy  the  apartment  we  were  sharing. 
The  date  was  set  for  Octotier  3rd  and  1  was 
honored  in  being  chosen  as  his  best  man  at  their 
wedding. 

"Goodrich  retired  in  1944  and  Henry  was  ap- 
pointed Curator  of  Mollusks.  He  inherited  a  divi- 
sion rich  in  historical  importance,  perhaps  the 
finest  collection  of  land  and  freshwater  mollusks 
in  the  country,  and  a  top  notch  library.  Unfor- 
tunately the  yearly  budget  had  i)een  reduced  to 
poverty  level.  Henry  didn't  remain  discouraged 
for  very  long,  however,  and  his  frugality,  cou- 
pled with  his  exploring  abilities  that  might  pro- 
vide increments  to  purchase  microsco[)es, 
microtomes,  establishing  aquarium  rooms,  etc., 
not  only  restored  the  budget  to  its  original 
status,  but  surpassed  it  many  times.  The  new 
addition  to  the  Museums  Building  was  con- 
structed and  this  allowed  the  Mollusk  Division  to 
expand  in  scientific  areas  uniciue  in  malacology. 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS     119 


The  Division's  liigh  status  subsequently  at- 
tracted splendid  investigators  to  receive  their 
advanced  training  in  malacology  at  the  Univer- 
sity of  Michigan  under  Henry's  direction. 

"I  well  recall  two  phrases  Henry  frequently 
used,  particularly  when  it  involved  a  person  with 
whom  he  disagreed.  He  would  say,  "Well  let  me 
tell  you,  I've  got  news  for  him!"  This  remark 
would  be  followed  with  what  he  was  going  to  do, 
and  he  wasn't  fooling!  The  magnificent  achieve- 
ments accomplished  by  the  Mollusk  Division 
were  due  to  Henry's  tremendous  effort  and 
unbelievable  drive.  When  deeply  disappointed 
by  a  granting  agency's  reply  to  a  grant  request 
which  Henry  had  painstakingly  made,  and  which 
had  looked  promising,  that  stated,  'We  regret  to 
inform  you.  .  .'  Henry,  having  given  vent  to  his 
feelings,  would  end  the  discussion  with  a  grin 
and  a  shrug  of  his  shoulders,  saying,  'Oh  well, 
we're  having  fun!'  " 

He  meant  it!  An  inspiring  teacher  and  fierce 
advocate  for  rigorous  study  and  understanding 
of  mollusks,  Henry  van  der  Schalie  expected  a 
lot  from  us.  He  expected  us  to  be  rational, 
careful,  committed,  and  to  derive  joy  from  our 
dedication,  as  he  did  from  his.  As  one  of  his  last 
students,  I  have  treasured  the  great  legacy  of 
his  intellect  and  his  spirit.  I  never  knew  "Van"  to 
be  half-hearted  about  anything.  He  was  em- 
phatic, enthusiastic  about  whatever  he  was  do- 
ing, whether  it  was  digging  into  a  new  research 
project,  encouraging  a  student,  pursuing  his 
beloved  mollusks  in  their  improbable  habitats, 
or  sharing  his  parenting  energies  with  his  in- 
comparable wife  and  colleague,  Annette.  He 
regularly  doted  on  all  the  accomplishments  of 
his  children,  each  of  whom  he  and  Annette 
always  regarded  with  boundless,  affectionate 
concern  and  pride. 

Van  had  a  strong  environmental  ethic  long 
before  the  raised  consciousness  of  the  present 
era.  He  evinced  articulate,  righteous  indigna- 
tion whenever  he  perceived  that  human  error, 
indifference  or  greed  had  resulted  in  the 
destruction  of  a  precious  habitat,  or  failed  to 
fund  research,  or  allowed  neglect  of  irreplace- 
able museum  collections.  I  remember  his  re- 
counting to   me  his  participation   in   a  panel 


discussion,  along  with  other  biologists  and 
lawyers,  on  the  University  of  Michigan  campus, 
in  the  1970's,  on  aspects  of  environmental  law. 
Van  related  to  me  his  consternation  with  the  at- 
titude evinced  by  his  colleagues,  who  had  caved 
in  to  sophistry  and  had  adopted  a  resigned, 
expedient  demeanor  in  the  discussion.  Van  ex- 
postulated, "My  God!  When  I  think  of  what 
(natural  resources)  we  have  squandered!"  Van 
insisted,  rightly  I  think,  that  better  laws  to  pro- 
tect the  environment  could  not  be  made  in  a 
society  where  knowledgeable  experts  didn't 
care.  He  believed  passionately  in  the  coupling  of 
the  fine  old  values  of  knowledge  and  respon- 
sibility. Van's  capacity  for  moral  outrage  was 
good  for  us  all.  It  continues  to  inspire  and  sus- 
tain us  as  we  pursue  urgent,  sometimes  un- 
popular, environmental  causes  of  our  own. 

In  recent  years,  a  visit  to  Manchester, 
Michigan,  to  see  Van  and  Annette  for  good 
mollusk  talk,  in  their  palatial  spread  along  the 
Raisin  River,  was  a  revitalizing  joy.  One  ambled 
with  them  about  their  acres,  past  the  huge, 
burgeoning  vegetable  gardens  Van  and  Annette 
and  their  son,  Arnold,  were  cultivating  in 
delighted  competition  with  each  other.  One 
followed  them  through  the  verdant,  leafy 
marshland  next  to  the  river,  where  Van  would 
pridefully  point  out  the  most  recent  wonders  of 
Annette's  botanical  expertise,  and  then  to  the 
Raisin  River  itself,  where,  thanks  to  van  der 
Schalie  effort,  a  number  of  species  of  Lampsilis 
still  maintain  reproducing  populations. 

On  such  jaunts  I  always  got  a  lump  in  my 
throat.  I  knew  again  that  this  wonderful  man, 
Henry  van  der  Schalie,  lived  a  life  of  indefatig- 
able personal  integrity.  His  career  spanned 
almost  five  decades  of  vital  research  at  the 
University  of  Michigan  and  around  the  world. 
More,  to  the  end,  he  cared  deeply  about 
mollusks  as  mollusks,  about  human  beings  as 
human  beings.  In  a  glowing,  over-arching  way 
Van  cared  about  the  lovely  planet  that  we  share. 
Such  jaunts  invariably  left  me  with  the  poignant 
reminder  that  I  shall  always  think  of  Van  as  a 
grand  model.  I  shall  always  think  of  Van  with 
admiration  and  love. 


120    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


CANTHARUS  MULTANGULUS  NEW  SUBSPECIES 
GRANDANUS  FROM  NORTHWEST  FLORIDA  (BUCCINIDAE) 


R.  Tucker  Abbott 

American  Malacologists,  Inc. 
Melbourne,  Florida  32902-2255 


During  the  last  few  years,  members  of  the 
Gulf  Coast  Shell  Club  in  Panama  City,  north- 
west Florida,  have  been  very  active  in  surveying 
the  marine  mollusks  of  their  area.  In  fact,  they 
have  published  several  checklists  and  in  1983 
issued  their  illustrated  Seashells  of  Bay  County 
and  the  Gulf  Coast.  In  1982,  Robert  Granda,  and 
later  other  members,  collected  and  sent  me  an 
unusual  form  of  Cantharus  which  I  now  take 
pleasure  in  naming  after  its  original  discoverer. 

Family  Buccinidae  Rafinesque,  1815 
Genus  Canthnrn.'^  Roding,  1798 
Cantharus  tnultangulus  (Philippi,  1848) 
new  subspecies  grandanus 
Descrij)tiom.-?>\\Q\\   small,   up   to   28   mm   in 
length,  fusiform,  light-weight,  finely  but  coarse- 
ly sculptured,  and  with  6  or  7  whorls.  Nuclear 
whorls  IV2,  smooth,  rounded,  rapidly  descend- 
ing,   translucent   tan    to   clear.    First   4   or   5 
postnuclear  whorls  slightly  shouldered,  with  10 
or  11  strong,  rounded,  axial  ribs  crossed  by 


FIG.  1.  Canlhariix  muUanifuiu.s  subspecies  ynmdanux 
Abbott,  holotype,  UNNM  no.  8.59096,  28.7  x  14.4  mm.  FIG. 
2.  Cantharus  multangulus  multangulus  (F^hilippi,  1848) 
from  off  Shell  Island,  Panama  City,  N.W.  Florida,  29.1  x 
15.6  mm.  Both  collected  by  Robert  Granda,  1982. 


about  9  or  10  very  small,  irregularly-sized, 
square-topped,  spiral  threads.  On  the  penulti- 
mate and  last  whorl,  which  is  smoothly  rounded, 
the  axial  ribs  are  obsolete.  Body  whorl  and  short 
siphonal  canal  with  about  30  to  40  fine  spiral 
threads  of  uneven  size.  Columella  with  two 
weak,  oblique  plicae  at  the  base.  Outer  lip  simple 
and  minutely  jagged.  Inside  of  last  whorl  glossy 
tan  and  with  about  a  dozen,  smooth  spiral 
threads.  Color  of  outer  shell  tannish  brown,  red- 
dish or  whitish  yellow  with  numerous  fine 
flecks,  occasional  narrow  spiral  bands  and 
sparse  axial  flames  of  chocolate-brown.  Oper- 
culum chitinous,  translucent  tan,  elongate-oval 
and  filling  most  of  the  aperture.  Soft  parts  and 
radulae  not  examined. 

Type  locality -2  to  4  feet  of  water,  on  sand 
and  weed  bottom.  Islet,  V4  mi  east  of  Black  Isle, 
Saint  Joseph  Bay,  off  Port  St.  Joe,  Gulf  County, 
northwest  Florida.  Robert  Granda,  collector, 
March  13,  1982. 

Types -The  holotype  is  deposited  in  U.  S.  Na- 
tional Museum  as  no.  859096.  Parat>']oes  from 
the  type  locality  have  been  deposited  in  the 
Academy  of  Natural  Sciences  of  Philadelphia, 
the  American  Museum  of  Natural  History,  N.Y., 
the  Museum  of  Comparative  Zoology  at  Har- 
vard, and  the  Florida  State  Museum,  University 
of  Gainesville,  Florida. 

Records -This  moderately  rare  subspecies  has 
been  collected  in  several  places  in  St.  Joe  Bay. 
Linda  Brunner  has  also  collected  this  subspecies 
in  St.  Andrews  Bay  in  nearby  Bay  County. 

Measurements  (mm.)- 


length 

width 

Holotype 

28.7 

14.4 

F'aratype 

26.8 

13.1 

Paratype 

26.7 

14.0 

i'aralype 

24.0 

12.1 

Paratype 

21.1 

11.0 

Paratype 

21.0 

11.0 

Remarks-The  subspecies  grandanus  of  the 
northeast  Gulf  of  Mexico  differs  from  the 
nominate    Cantharus    multangulus    (Philippi, 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    121 


1848)  in  having  a  lighter-weight  shell,  lacking 
the  strong  shoulder  nodules  on  the  last  two 
whorls,  in  having  more  numerous  spiral  threads, 
and  in  being  a  little  more  elongate  in  propor- 
tions. The  variations  in  colors  and  patterns  are 
very  similar.  It  is  possible  that  these  colonies  in 
the  "Panhandle"  region  of  Florida  represent  an 
ecological,  rather  than  a  genetic,  form. 

Mr.  Granda  obtained  two  "clutches"  of  small, 
horny  egg-capsules  which  he  found  on  pieces  of 
carapace  from  the  horseshoe  crab,  Limulus.  The 
urn-shaped  capsules,  about  5x8  mm,  closely 
resembled  those  so  well  illustrated  by  D'Asaro  in 
his  account  of  the  capsules  of  CantharuH 
multangulus  from  the  same  region  (1986,  p.  86, 
figs.  A-D).  Very  similar  capsules  of  the 
nominate  species  from  Sanibel  Island  were  il- 
lustrated by  Perry  and  Schwengel,  1955,  pi.  50, 
fig.  340. 

Cantharus  cancellarius  (Conrad,  1846)  from 
the  same  region  differs  in  lieing  more  ovoid, 
having  a  shorter  spire  and   in  having  much 


stronger  and  fewer  spiral  threads.  The  similar 
muricid,  Cnlotrophon  ostrearum  {Conrad,  1846), 
has  stronger  shoulder  nodes  and  a  mauve  to 
rosy-purple  aperture.  Fossil  C.  multangulus 
from  the  old  St.  Petersburg  pits  have  fewer 
axial  nodes  per  whorl.  This  grou])  of  species  ap- 
pears to  t>e  largely  confined  to  southeast  United 
States,  the  Bahamas,  the  north  coast  of  Cuba 
and  Yucatan,  Mexico. 

LITERATURE  CITED 

D'Asaro.  Charles  N.  1986.  Egg  Capsules  of  Eleven  Marine 

Prosobranchs  from  Northwest  Florida.  BtiU.  Murine  Sri. 

39(1):  76-91,  4  figs. 
Gulf  Coast   Shell   Club  (Bob   Cranda   and   -Jim   Brunner, 

editors).  1983.  SeaskeLls  of  Bay  Cininlii  mtd  the  Gi(lfC(t(it;t. 

26  pp.,  96  photos,  1  map. 
Perry,  Louise  M.  and  Jeanne  S.  Schwengel  19.5.5.  Murine 

Shells  of  the  Westeiyt  Coast  of  FUiridti.  Paleont.  Research 

Inst..  Ithaca.  318  pp,  55  pis. 
Robertson,  Robert.  1957.  A  Study  of  Cantharus  multaiigu- 

his    (Philippi)    with    Notes   on    Cantharus    and   Pseudo- 

neptunea    (Gastropoda:Buccinidae).    Notulae    Naturae. 

Philadelphia,  no.  300,  pp.  1-10,  19  figs. 


ON  THE  TAXONOMICAL  STATUS  OF  TRITONIUM  VIRIDULUM 
FABRICIUS,  1780  (GASTROPODA:  CANCELLARIIDAE) 

Jon-Arne  Sneli  and  Oystein  Stokland 

Trondhjem  Biologiske  Stasjon 
7000  Trondheim,  Norway 

ABSTRACT 

The  taxonoviic  and  nomenclatorial  problems  associated  with  the  species 
Admete  viridula  (Fahricius.  1780)  are  discussed.  The  fact  that  the  type  specimens 
of  both  Admete  viridula  and  A.  crisps.  Moller,  1842  are  misingfrom  the  Zoological 
Museum  in  Copenhagen  complicates  the  situation  considerably  as  three  species 
are  involved,  Tritonium  viridulum  Fabricius,  1780,  Defrancia  viridula  Moller, 
1842.  and  D.  exarata  Moller.  1842.  Fabricius's  name  is  to  be  regarded  as  a  nomen 
dubium  and  that  Admete  couthouyi  (Jay,  1839)  should  be  used  as  the  correct  name 
for  that  species.  Mailer's  D.  viridula  and  D.  exarata  are  both  good  species;  but  D. 
viridula  should  probably  be  named  Oenopota  decussata  (Couthouy,  1839)  and  D. 
exarata,  Propebela  exarata  (Moller,  1842). 


The  original  description  of  Tritonium 
viridulum,  which  was  published  by  Fabricius  in 
1780  without  an  illustration,  has  commonly  been 
regarded  as  the  original  description  of  Admete 
viridula  auctt.  The  description  fits  the  species 
fairly  well,  and  the  taxonomic  situation  would 


appear  quite  simple. 

However,  Ball  (1886:  298)  after  having  ex- 
amined the  holotype  in  the  Zoological  Museum 
in  Copenhagen,  claimed  that  the  species  be- 
longed to  the  group  Bela  auctt.  He  also  found 
that  Defrancia  viridula  Moller,   1842  (Fig.   1) 


122    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


0H:- 


5  6  ^7 

FIGS.  1-7.  1,  Pro-pehela  exarata  (Moller,  1842);  one  of  the  larger  specimens  from  the  syntype-collection  of  Moller  (Zool.  nius., 
Copenhagen).  2,  Propebela  exarntri  (Moller)  as  drawn  by  G.  O.  Sars.  1878.  3  and  4,  A  specimen  of  Oenuputn  deccusata 
(Couthouy,  1839)  =  Bda  viriduta.  from  the  collection  of  Moller  (Zool.  mus,  Copenhagen).  On  the  label  is  also  written:  "B. 
viridula  (et  var.  inflata)  =  B.  deccusata  Couthouy  var.  ventricosa".  5,  Admete  viriduta  uctt.  as  drawn  by  G.  0.  Sars,  1878.  6 
and  7,  Holotype  of  Cancellaria  buccinoides  Couthouy,  1838  (Reg.  no.  279394  in  Mus.  Comp.  Zool,  Cambridge,  Mass.).  The 
species  is  Admete  couthouyi  Jay,  1839. 


was  founded  on  the  same  specimen,  and  con- 
cluded that  it  was  identical  with  Defrancia  ex- 
arata Moller,  1842  (Fig.  2).  Ball  writes  that 
Morch  had  discovered  these  facts  before  him 
and  Posselt  (1898:  168)  confirms  this  by  referr- 
ing to  a  handwritten  catalogue  by  Morch.  Con- 
cerning the  nomenclatorial  situation,  Posselt 
claims  that  the  correct  name  o^  Admete  viridula 
auctt.  (Fig.  3)  should  be  Admete  couthouyi  Jay, 
1839,  because  the  older  Cancellaria  buccinoides^ 
(Fig.  4)  of  Couthouy,  1838  was  described  in 
another  genus. 

All  three  species  involved  in  this  problem 
show  a  great  deal  of  variability,  and  Fabricius 
description  fits  both  Moller's  Defrancia  viridula 
and  Admete  viridula  auctt.  as  well.  Concerning 
Defrancia  exarata,  Posselt  (1898:  168)  states 
that  Admete  crispa  Moller  1842,  which  conmion- 


ly  is  regarded  as  conspecific  with  A.  viridula 
auctt.,  resembles  D.  exarata  with  respect  to 
sculpture. 

This  situation  could  have  easily  been  cleared 
up  if  the  type  specimens  in  question,  which  were 
all  from  Greenland,  had  been  available,  but  un- 
fortunately the  type  specimens  of  both 
Tritonium  riridulum  and  Admete  crispa  appear 
to  have  disappeared  early  in  this  century 
(Jorgen  Knudsen,  pers.  commn.).  Only  one  sam- 
ple in  the  collection  of  the  Zoological  Museum  in 
Copenhagen  could  possibly  be  Fabricius  type, 
but  this  sample  contains  three  specimens 
whereas  the  original  description  states  that 
Fabricius  had  only  one  specimen  available.  In 
the  Zoological  Museum  in  Copenhagen  there  are 
type  lots  of  both  Mollers  species  Defrancia  ex- 
arata (Fig.  2)  and  D.  viridula  (Fig.  1).  The  type 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    123 


lots  of  the  last  species  contains  seven  syntypes 
and  thereby  shows  that  Moller  did  not  base  his 
description  on  Fabricius  specimen. 

None  of  Mollers  type  specimen  fits  very 
well  with  Fabricius  description.  However, 
Knipowitsch  (1901)  illustrates  a  specimen  oi  D. 
exarata  from  Svalbard  which  rather  closely 
resembles  Admete  viridula  auctt.  Considering 
this,  and  that  the  form  caWed  Admete  critipa  has 
a  sculpture  like  that  of  D.  exarata,  we  assume 
that  some  morphological  overlap  between  the 
species  is  present.  Although  fitting  the  common 
forms  of  Admete  viridula  auctt.  better  than 
those  of  Holler's  Dejrancia  exarata,  Fabricius 
description  covers  the  overlapping  forms,  with 
the  exception  that  we  never  have  observed  D. 
exarata  without  prominent  ribs  on  the  last 
whorl  as  mentioned  in  the  description.  How- 
ever, considering  the  large  intraspecific  varia- 
tion within  most  species  in  this  group  it  is  no 
wonder  that  Dall  (1886)  reduced  them  to 
synonomy. 

Moller's  sample  of  Defrancia  viridula  fits 
Fabricius  description  fairly  well  except  that  the 
ribs,  which  are  said  to  be  straight  ("costae 
longitudinales")  in  Fabricius  description,  are 
somewhat  curved.  As  Moller's  Defrancia 
viridula,  which  probably  is  conspecific  with 
Pleurot.oma  decussata.  Couthouy,  1839,  has  a 
large  intraspecific  variation  like  all  species  in 
this  group,  Fabricius  description  fits  both 
species  fairly  well,  and  one  cannot  apply  it  to 
one  rather  than  the  other. 

Concerning  Morch's  earlier  opinions  as  first 
published  by  Dall  (1886),  there  is  a  sample  in  the 
Zoological  Museum,  University  of  Bergen  (no. 
28208)  which  should  be  mentioned.  This  con- 
tains two  dry  specimens  of  Moller's  viridula 
from  Greenland  which  are  identified  to  "Bela 
viridula  M.  Sars"  by  Morch.  Michael  Sars  never 
described  any  species  with  this  name,  but  the 
sample  could  indicate  that  Morch  had  Moller's 
species  in  mind  when  he  claimed  to  have  dis- 
covered that  this  and  Fabricius  species  was 
founded  on  the  same  specimen.  However,  this  is 
in  contradiction  to  the  presence  of  the  seven 
syntypes  of  Defrancia  viridula  Moller  in  the 
Zoological  Museum  of  Copenhagen. 

Some  additional  information  about  the  prob- 
lems within  the  genus  Admete  may  be  found  in 
Troschel's    (1866-1893)    work    on    gastropod 


radulae.  Troschel  investigated  preserved 
material  of  both  Defrancia  viridula  and  the 
genus  Admete  in  the  Zoological  Museum  in 
Copenhagen  with  respect  to  radular  teeth.  He 
found  the  radulae  in  D.  viridula  were  like  those 
of  other  species  within  Beta  auctt.,  while  in 
Admete  he  found  two  kinds  of  teeth.  He  also 
found  two  shell  forms  which  he  called  Adm,ete 
viridula  (Fabricius)  and  A.  crispa  Moller  accor- 
ding to  figures  in  Middendorffs  (1849)  work  on 
Russian  mollusks.  The  shell  forms  corresponded 
with  the  radular  forms,  which  may  indicate  that 
Troschel  was  correct  in  dividing  the  traditional 
Admete  viridula  into  two  species. 

The  nomenclatorial  consequences  of  these  tax- 
onomic  circumstances  may  now  be  summarized. 
Moller's  Defrancia  exarata  presents  no  prob- 
lems, and  in  our  opinion  it  is  best  placed  within 
Propebela  Iredale,  1918.  Fabricius  Tritionium 
viridulum  seems  difficult  to  identify  with  any 
species,  and  in  our  opinion  it  should,  since  the 
holotype  is  lost,  be  regarded  as  a  nomen  dubium. 
One  could  use  the  existence  of  the  sample  iden- 
tified by  Morch  to  attach  the  name  to  Moller's 
viridula,  having  also  the  facts  presented  by  Dall 
in  mind.  However,  since  both  Morch  and  Dall 
seems  to  have  considered  that  Fabricius  original 
specimen  belonged  to  Propebela  exarata  (Moller, 
1842),  this  argument  seems  somewhat  dubious. 
The  fact  that  Moller  founded  his  Defrancia 
viridula  on  a  sample  of  seven  syntypes  still  pre- 
sent in  the  Zoological  Museum  in  Copenhagen 
also  contradicts  the  view  of  Morch  and  Dall.  In 
our  opinion,  Moller's  Defrancia  viridula  is  best 
placed  in  Oenopota  Morch,  1852,  probably  as  a 
synonym  of  Oenopota  decussata  (Couthouy, 
1839). 

Concerning  ylrf?nete  viridula  auctt.,  the  oldest 
name  for  this  species  is  Cancellaria  buccinoides 
Couthouy,  1838.  This  name  is,  however,  a 
primary  homonym  of  Cancellaria  buccinoides 
W.  Wood,  1828  and  has  to  be  rejected  as  there 
are  no  strong  reasons  why  it  should  be  referred 
to  the  International  Commission  on  Zoological 
Nomenclature  for  eventual  preservation.  The 
next  available  name  is  Admete  couthouyi  Jay, 
1839,  which  meets  all  demands  as  a  correct 
name  for  the  species.  The  type  of  this  is  also 
Couthouy 's  type  of  buccinoides  since  Jay's  name 
was  proposed  as  a  nomen  novum.  An  eventual 
separation  of  Mollers  Adynete  crispa  from  A. 


124    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


couthouyi  as  proposed  by  Troschel  (1866-1893) 
will  not  be  discussed  in  detail,  but  as  the  type 
specimen  of  A.  crispn  is  lost  either  a  neotype 
should  be  selected  or  a  new  name  should  be  pro- 
posed and  Mdller's  name,  A.  crispa,  regarded  as 
a  nornen  dubium.  As  MoUer  never  described  the 
radula  of  his  species  the  second  possibility  is 
probably  the  best. 

Mr.   Georg  Crawford   kindly   corrected   our 
English  text. 

LITERATURE  CITED 

Couthouy,  J.  P.  1838.  Descriptions  of  new  species  of 
Mollusca  and  shells  and  remarks  on  several  polypi  found  in 
Massachusetts  Bay.  Boston  Jour.  Nat.  Hist.  2:53-111. 

1839.  Monograph  on  the  family  Osteodesmacea 

of  Deshayes,  with  remarks  on  two  species  o(  Patelloidea, 
and  descriptions  of  new  species  of  marine  shells,  a  species 
of  Anculotu.'i.  and  one  of  Eolis.  Boston  Jour.  Hist. 
2:129-189. 

Dall,  W.  H.  1886.  Supplementary  notes  on  some  species  of 
mollusks  of  the  Bering  Sea  and  vicinity.  Proc.  U.  S.  natn. 
M«,s.  9:297-309. 


Fahricius,  0.  1780.  Faumi  Groenliindini.  Johannes  Gottloli 

Rolhe,  Kobenhavn.  452  pp. 
Iredale,  T.   1918.  Molluscan  nomenclatural  prulilems  and 

solutions. -No.  1.  Proc.  Malac.  Soc.  Land.  13:28-40. 
Jay,  J.  C.  1839.  A  catalogue  of  shells  in  the  collection  of  John 

C.  Jay,  M.D.,  with  descriptions.  Ed.  3.  125  pp. 
Knipowitsch,  N.  1901.  Ueber  die  in  den  Jahren  1899-1900 

im  Gebiete  von  Spitzbergen  gesammelten  recenten  Mollus- 

ken  und  Brachiopoden.  Ezheg.  zool.  Muz.  6:435-558. 
Middendorff,  A.  T.  v.  1849.  Beitrage  zu  einer  Malacozoologia 

Rossica.  Abt.  2  &  3.  Mem.  Acad.  Imper.  Sci.  St.-Petersb.. 

ser.  6.  Sci.  nat.  6:1-187. 
Moller,  H.  P.  C.  1842.  Index  Molluscorum  Groenlandiae. 

Naturhist.  Tids.'^kr.  ser.  1.  4:76-97. 
Posselt,    H.    J.    (A.    S.    Jensen    ex.)    1898.    Grondlands 

brachiopoder  og  bloddyr.  Meddr.  Gronland  23:1-298. 
Sars,   G.   O.    1878.   Mollusca  regionis  arcticae  norvegiae. 

Bidrag  til  kundskaben  oni  Norges  nrkliske  fiiuna.  A.  W. 

Brogger,  Christiania  (Oslo).  446  pp. 
Trochel.  F.  H.  1866-1893  (cont.  by  J.  Thiele  in  1891).  Das 

Gebiss  der  Schnecken  zur  Begmndung  einer  naturlichen 

Classification.   Vol.   2.   Nicolaische   Verlags-Buchhandl., 

Berlin. 
Wood,  W.  1828.  Index  Testnceoi.Ed.  2.  with  the  plates  and 

suppL,  London.  188  |)p. 


PISIDIUM  HENSLOWANUM  (SHEPPARD)  IN  THE 
CONNECTICUT  RIVER,  MASSACHUSETTS  (BIVALVIA:  PISIDIIDAE) 

Douglas  G.  Smith 

Museum  of  Zoology 

University  of  Massachusetts 

Amherst,  Massachusetts  01003-0027 


ABSTRACT 
The  pisidiid  dam  Pisidium  henslowanum  ha^  been  coyisidered  by  several 
authors  as  an  introdnced  species  in  North  America  despite  its  discovery  in 
several  parts  of  North  America  well  away  from  industrial  centers.  The  species 
has  been  recently  found  in  the  Connecticut  River  in  Massachusetts  which  is  the 
southeastern  most  record  yet  in  North  America.  It  is  suggested  that  this  .species  is 
native  but  locally  distributed.  The  Connecticut  River  record  seems  also  to  repre- 
sent the  softest  water  in  which  the  species  has  been  reported  in  North  America. 


Since  its  first  discovery  in  North  America  by 
Sterki  (1899),  the  pisidiid  clam,  Pisidium 
henslowanum  (Sheppard,  182.'j),  was  known  on 
the  continent  from  only  a  small  number  of  sites 
mostly  clustered  in  the  Great  Lakes  and  St. 
Lawrence  River  drainages  (Heard,  1961,  1962; 
Ilerrington,  1962).  The  species  is  well  known  in 
the  palearctic  region  (Woodward,  1913;  Zhadin, 


1957;  Ellis,  1978).  Herrington  (1962,  1965)  and 
Heard  (1962)  determined  that  P.  henslowanum 
was  introduced  into  North  America.  Subsequent 
to  Herrington's  (1962)  and  La  Roccjue's  (1967) 
reviews  of  the  North  American  distribution  of 
P.  henslou'anum.  the  species  was  reported  from 
a  number  of  localities  in  central  and  western 
Canada  by  Harris  (1973),  who  also  documented 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    125 


the  occurrence  of  P.  henslowanum  in  7000  year 
old  deposits.  Harris  (1973)  used  this  information 
to  question  Herrington's  (1962)  and  Heard's 
(1962)  conclusions  that  the  species  was  not 
native  to  North  America.  Harris  (1973)  also 
cited  Herrington's  (1962)  listing  of  a  much 
earlier  fossil  date  (Pliocene-Pleistocene)  for  P. 
supinum,  then  considered  by  Herrington  a 
synonym  of  P.  henslowanum.  However,  subse- 
quent study  led  Herrington  (1965)  to  accept  P. 
supinum  as  a  distinct  species,  thus  weakening 
Harris'  (1973)  argument.  Nonetheless,  Harris' 
(1973)  data  compelled  a  reassessment  of  the  in- 
troduced status  of  P.  henslowanum,  yet  subse- 
quent studies  (Burch,  1975;  Clarke,  1981; 
Mackie,  1981)  continued  to  follow  Herrington 
(1962,  1965)  and  Heard  (1962). 

During  the  summer  of  1985,  P.  henslowanmn 
was  collected  in  the  Connecticut  River  at  two 
locations  in  Hampden  County,  Massachusetts. 
The  first  collection  was  made  in  the  Holyoke  in- 
dustrial canal  complex  (in  direct  communication 
with  the  river)  in  Holyoke  and  the  second  in  the 
river  in  Longmeadow.  The  Holyoke  collections 
were  first  made  on  1  July  and  repeated  on  27 
October.  Prior  to  each  collection  the  canals  had 
been  drained  for  routine  maintenance.  A  total  of 
98  specimens  were  collected,  of  which  a  series  of 
shells  have  been  deposited  into  the  Museum  of 
Comparative  Zoology,  Harvard  University, 
Cambridge,  Massachusetts  (MCZ).  The  remain- 
ing alcohol  preserved  material  has  been  placed 
into  the  Museum  of  Zoology,  University  of 
Massachusetts  at  Amherst.  The  Longmeadow 
collection  was  made  on  16  September  and  com- 
prised a  single  adult  specimen.  Previous  to  these 
collections  the  only  New  England  records  for  P. 
henslowanum  were  Lake  Champlain  in  Vermont 
(St.  Lawrence  River  system)  and  Aroostook 
County,  Maine  (Johnson,  1915).  Although  the 
Vermont  record  is  generally  accepted  in  subse- 
quent literature,  the  Maine  record  has  not  been 
repeated.  The  Maine  record  was  listed  by 
Lermond  (1909)  as  well  and  was  credited  to 
Nylander  as  collector  and  Sterki  (presumably) 
as  the  authority.  A  search  of  the  pisidiid  collec- 
tions at  the  MCZ  revealed  no  specimens  of  P. 
henslowanum  from  either  locality. 

The  Connecticut  River  is  clearly  outside  the 
Great  Lakes  and  St.  Lawrence  River  drainages 
and  the  populations  in  the  Connecticut  River 


could  not  have  been  established  by  recent  migra- 
tion. Additionally,  the  section  of  the  Connec- 
ticut River  in  Massachusetts  is  well  above  the 
commercially  navigable  portion  of  the  river  and 
it  seems  unlikely  that  the  population  in  the 
Holyoke  canals  has  been  artificially  founded  by 
"stowaways"  from  Europe.  As  argued  by  Harris 
(1973),  this  species  may  not  be  introduced  in 
North  America.  It  might,  however,  be  native 
but  very  localized  and  thereby  easily  overlooked. 
The  Holyoke  canal  collections  were  made  in  mud 
and  silt  sediments  normally  under  4.5  m  of 
water.  The  single  Longmeadow  specimen  was 
collected  in  .5  m  of  water.  The  species  is  com- 
mon in  the  Holyoke  canals,  particularly  along 
the  base  of  the  retaining  walls,  and  is  associated 
with  the  pisidiid  species  Musculium  securis  and 
Sphaerium.  striatinum  and  juveniles  of  the 
unionid  mussels  Elliptio  complanata  and  Ano- 
donta  implicata.  Chemical  data  for  the  canals 
(Anon.,  1978,  1980)  includes  the  following  para- 
meters (during  July,  August  and  September): 
pH  =  7.0-7.7,  total  alkalinity  =  25-35  mg/1 
(CaC03),  dissolved  oxygen  =  5.2-10.4  mg/1, 
nitrate  =  0.0-0.5  mg/1,  chloride  =  7-35  mg/1. 
The  chloride  values  in  part  represent  runoff  of 
residues  of  municipal  operations.  Chlorides  in 
the  Longmeadow  portion  of  the  river  average 
about  10  mg/1  (Anon.,  1978).  These  values  com- 
pare well  with  data  given  by  Harris  (1973)  and 
Okland  and  Kuiper  (1980)  except  for  the  alkalin- 
ity values.  In  Europe,  Ellis  (1978)  has  indicated 
that  P.  henslowanum  prefers  calcium  rich 
waters  and  Harris  (1973)  concludes  that  the 
presence  of  dissolved  salts  are  an  important 
characteristic  of  this  species'  habitat.  Overall, 
the  water  of  the  Connecticut  River  is  relatively 
soft  and  has  average  levels  of  chloride  salts 
(Wetzel,  1975).  Thus  P.  henslowanum  can  ap- 
parently exist  in  somewhat  soft  water. 

Specimens  collected  on  1  July  were  for  the 
most  part  larviparous  (11  of  13  specimens  or 
84%  of  the  sample),  and  the  16  September 
specimen  from  Longmeadow  was  larviparous 
also.  Of  the  27  October  sample,  however,  only 
2%  (2  of  85)  were  larviparous  though  many  con- 
tained egg  masses  in  the  gills.  These  seasonal 
data  compare  well  with  Holopainen  and 
Jonasson's  (1983)  findings  for  the  presence  of 
brooded  larvae  in  Danish  populations. 


126    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


Acknowledgments 

I  thank  Alan  Richmond  and  Jenifer  Bush  for 
assistance  in  the  field.  I  also  thank  D.  Kenneth 
Boss  for  allowing  access  to  collections  at  the 
MCZ,  Harvard  University. 

LITERATURE  CITED 

Anonymous.  1978.  The  Connecticut  River  water  quality 
data- 1978.  Massachusetts  Department  Environmental 
Quality  Engineering.  Division  Water  Pollution  Control. 
Westborough,  Massachusetts.  147  pp. 

.  1980.  Connecticut  River  Water  Quality  Data- 

1980.  Massachusetts  Department  Environmental  Quality 
Engineering.  Division  Water  Pollution  Control.  West- 
borough,  Massachusetts.  60  pp. 

Burch,  J.  B.  1975.  Freshwater  Sphaeriacean  Clams  (Mol- 
lusca:  Pelecypoda)  of  North  America.  Malacological  Pub- 
lications, Hamburg,  Michigan.  96  pp. 

Clarke,  A.  H.  1981.  The  Freshivaler  Molluscs  of  Canada. 
National  Museums  of  Canada,  Ottawa.  446  pp. 

Ellis,  A.  E.  1978.  British  Freshwater  Bivalve  Mollusca. 
Synopsis  of  the  British  Fauna  (New  Series)  No.  11. 
Academic  Press,  London.  109  pp. 

Harris,  S.  A.  1973.  Pisidium  henslowanum  (Sheppard)  in 
western  Canada.  The  Nautilus  87:86-87. 

Heard,  W.  H.  1961.  Pisidium  henslouianum  (Sheppard)  in 
Lake  Michigan.  The  Nautilu.^  74:12.3. 

1962.  Distribution  of  Sphaeriidae  (Pelecypoda) 

in  Michigan,  U.S.A.  Malacologia  1:139-161. 

Herrington,  H.  B.  1962.  A  revision  of  the  Sphaeriidae  of 
North    America   (Mollusca:    Pelecypoda).    Miscellaneous 


Publications  Museum  of  Zoology.  Univ.  Michigan.  No.  118. 
174  pp. 
1965.  Corrections  of  sphaeriid  nomenclature. 


The  Nautilus  79:42-45. 

Holopainen,  L  J.  and  P.  M.  Jonasson.  1983.  Long-term  popu- 
lation dynamics  and  production  of  Pisidium  (Bivalvia)  in 
the  profundal  of  Lake  Esrom,  Denmark.  Oikos  41:99-117. 

Johnson,  C.  W.  1915.  Fauna  of  New  England  13.  List  of  the 
Mollusca.  Occasional  Papers  Boston  Society  Natural 
History  7(13):1-231. 

Lermond,  N.  W.  1909.  Shells  of  Maine.  Report  of  the  Com- 
missioner of  Agriculture  of  Maine,  1909.  Pp.  217-262. 

Mackie,  G.  L.  1981.  Nearctic  freshwater  Sphaeriacea 
(Bivalvia).  Bulletin  Amfrican  Malacological  Union  for 
1981:  49-52. 

Okland,  K.  A.  and  J.  G.  J.  Kuiper.  1980.  Small  mussels 
(Sphaeriidae)  in  freshwater  in  Norway-distribution, 
ecology',  and  relation  to  acidification.  SNSF  project,  IR 
61/80.  Oslo-As,  Norway.  85  pp. 

Sterki,  V.  1899.  Pisidia  new  to  our  country,  and  new  species. 
The  Nautilus  13:9-12. 

Wetzel,  R.  G.  1975.  Limnology.  W.  B.  Saunders  Co.,  Phila- 
delphia. 743  pp. 

Woodward,  B.  B.  1913.  Catalogue  of  the  British  species  of 
Pisidium  (Recent  and  fossil)  in  the  collections  of  the 
British  Museum  (Natural  History),  with  notes  on  those  of 
western  Europe.  British  Museum  (Natural  History),  Lon- 
don. 144  pp. 

Zhadin,  V.  I.  1952.  MoUusks  of  fresh  and  brackish  waters  of 
the  U.S.S.R.  Zoological  Institute  Academy  Sciences 
U.S.S.R.  No.  46.  (1965  Israel  Program  Scientific  Transla- 
tion. 365  pp.) 


PRELIMINARY  OBSERVATIONS  OF  PREDATION  ON  OCEAN 
QUAHAUGS,  ARCTICA  ISLANDICA,  BY  ATLANTIC  WOLFFISH, 

ANARHICHAS  LUPUS 

C.  M.  Hawkins  and  R.  B.  Angus 

Fisheries  Research  Branch,  Scotia-Fundy  Region 

Halifax  Fisheries  Research  Laboratory 

Halifax,  Nova  Scotia  B3J  2S7 


ABSTRACT 
Populations  of  the  ocean  quahaug,  Arctica  islandica,  may  be  siisceptable  to  in- 
tensive grazing  pressure  by  the  Atlantic  wolffish.  Anarhichas  lupus.  In  stomach 
analysis  of  four  wolffish  from  an  inshore  Nova  Scotia  area,  85-99%  of  the  wet 
weight  contents  were  Arctica  islandica.  The  number  of  quahaugs  in  the  stomach 
contents  ranged  from  3-11.  These  individuals  were  35-86  mm.  in  shell  length  and 
were  4-60  years  old. 


The  ocean  quahaug  (or  quahog),  Arctica  is- 
landica (Linnaeus,  1767),  is  an  underexploited 
commercial  mollusk  found  in  coastal  waters  of 


the  south  shore  of  Nova  Scotia,  Canada,  and  on 
the  Scotian  Shelf  (Rowell  and  Chaisson,  1983). 
These  bivalves  are  slow*  growing  (Murawski  et 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    127 


al,  1982)  reaching  a  minimum  market  size  of 
about  51  mm  (2  inches)  in  13  years,  and  long- 
lived  (225  years,  Ropes  and  Murawaski,  1983). 
Since  they  are  commercially  important  in  the 
northeast  of  the  United  States  (Murawski  et  a/., 
1982)  and  are  of  potential  commercial  impor- 
tance in  the  south  shore  area  of  Nova  Scotia  it  is 
important  for  the  fishery  biologist  and  manager 
to  assess  the  impact  of  natural  predation  on 
stocks  in  relation  to  stock  assessment  and 
potential  yields.  Quantative  data  on  the  diet  of 
the  common  Atlantic  wolffish  is  sparce  as  are 
data  on  natural  predation  of  ocean  quahaugs. 
Early  works  (Verrill  1871,  Gill  1911  and  others) 
are  primarily  qualitative  but  indicate  that  shell- 
fish play  an  important  role  in  wolffish  nutrition. 
Bowman  et  al.  (1976)  provide  more  quantitative 
data.  They  examined  the  stomach  contents  of 
127  wolffish  from  offshore  waters  of  western 
Nova  Scotia.  Approximately  42%  of  all 
stomachs  were  empty,  but  the  remainder  on 
average  contained  about  28%  (by  wet  weight) 
bivalves.  However,  no  further  taxonomic  break- 
down was  made.  Although  there  is  evidence  of 
offshore  predation  of  Ardica  islandica  by  cod 
{Gadus  morhua)  (Arntz  1974,  1978)  there  is  a 
lack  of  both  qualitative  and  quantitative  data 
relating  to  predator  induced  natural  mortality 
ofArctica  islandica  in  coastal  waters.  From  our 
investigation  in  the  coastal  waters  of  Nova 
Scotia,  Canada,  qualitative  and  quantitative 
data  make  the  common  Atlantic  wolffish, 
Anarhichas  lupus,  a  prime  suspect  as  an  impor- 
tant predator  oi  Ardica  islandica. 

During  the  first  of  a  three  year  study  begun  in 
1982  to  study  maturation  oi  Ardica  islandica  at 
Port  Mouton,  Nova  Scotia  (64  5rN,  43  56'W) 
one  of  us  (RBA)  noted  the  arrival  of  wolffish  to 
the  study  site  in  the  spring  and  their  departure 
in  late  summer.  The  time  of  this  annual  migra- 
tion corresponded  to  wolffish  bycatches  in  net 
and  traps  reported  by  local  fishermen  operating 
in  other  areas  near  our  study  site.  The  fisher- 
men also  noted  that  the  wolffish  guts  were  full 
of  shell  material.  This  information  prompted  us 
to  catch  wolffish  at  our  study  site  to  determine 
whether  or  not  they  were  feeding  on  quahaugs 
and,  if  so,  their  impact  on  known  quahaug 
stocks. 

The  study  site,  about  4000  m^  of  flat  sandy 
bottom,    had    been    surveyed    30    times    with 


SCUBA  prior  to  sampling  for  wolffish.  The  area 
was  found  to  be  generally  void  of  any  macro- 
epibenthos  commonly  eaten  by  wolffish,  i.e. 
crabs,  gastropods,  and  sea  urchins  (Bigelow  and 
Schroeder,  1953).  Wolffish  inhabiting  this  area 
were  collected  by  a  diver  who  speared  the  fish 
and  returned  to  the  support  craft  where  fresh 
weight  {+/-  0.1  kg)  overall  length  (+/-  0.5  cm) 
and  sex  were  recorded.  Whole  intact  stomachs 
were  then  removed  and  fixed  in  10%  formalin  in 
seawater. 

In  the  laboratory  the  total  weight  of  the 
stomach  contents  (excluding  liquids)  was  first 
determined.  Subsequently,  stomach  contents 
were  sorted  and  recognizable  materials  were 
pooled  prior  to  weighing  (  +  /-  0.1  g  wet 
weight).  The  number  and  size  of  quahaugs  eaten 
by  wolffish  was  then  determined.  Recognizable 
umbo  portions  of  valves  were  separated  from 
stomach  contents  through  careful  examination 
of  shell  fragments.  The  number  of  paired 
umbones  was  considered  representative  of  the 
number  of  quahaugs  eaten.  The  original  size  of 
the  live  quahaugs  eaten  was  determined  by 
matching  the  fragmented  umbo  pieces  with 
other  valve  fragments  to  assemble  whole  or  par- 
tial valves.  Size  was  attained  by  superimposing 
on  the  whole  intact  valves  of  a  known  size.  Suc- 
cess in  assembling  the  valves  contributed  to  con- 
fidence in  determining  the  numbers  and  sizes  of 
the  quahaugs  consumed.  From  previous  re- 
search on  the  biology  of  Ardica  islandica  by 
Rowell  and  Chaisson  (1983),  relationships  were 
developed  for  animal  size  (overall  length  in  cm) 
to  age  and  whole  fresh  weight  to  equivalent 
meat  weight.  Estimates  of  the  total  fresh 
weight  as  well  as  equivalent  meat  weight  con- 
sumed by  each  fish  was  then  determined  from 
the  number  and  size  composition  of  quahaugs  in 
gut  contents. 

The  general,  solitary,  non-schooling  nature  of 
wolffish  (Bigelow  and  Schroeder,  1953)  is  sup- 
ported by  our  findings  that  few  wolffish  were 
caught  at  our  study  site.  We  encountered  wolf- 
fish in  only  four  of  twelve  SCUBA  surveys  over 
three  months  of  study  (June  1983;  May,  July 
1984).  In  addition,  we  noted  that  once  a  wolffish 
was  removed  another  occupied  the  former  resi- 
dence position  by  our  return.  This  may  indicate 
that  wolffish  occupy  and  possibly  defend  large 
territories  which  may  exceed  the  4000  m^  area 


128    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


of  our  study  site. 

Only  five  woiffish  were  caught  for  stomach 
content  analysis  (Table  1).  The  largest  individual 
measured  101.6  cm  in  overall  length  and 
weighed  12.0  kg,  the  smallest  was  72.6  cm  and 
4.4  kg  wet  weight.  Four  fish  contained  easily 
identifiable  quahaug  shells  (Arctica  islandica), 
comprising  85-99%  of  the  total  gut  content  wet 
weight  (Table  1).  The  remaining  foods  (1-15%) 
were  trace  amounts  of  periwinkles  {Littorina 
sp.)  sea  urchins  (Stronglyocentrotus  droe- 
hachiensis)  and  unidentifiable  crustacean  exo- 
skeleton.  The  fifth  fish  was  packed,  to  a  point 
where  the  stomach  was  distended,  with  finely 
crushed  tests  of  the  sanddollar  Echinarchinus 
parma. 

After  assembling  shell  fragments  for  qualita- 
tive analysis,  we  found  from  3-11  quahaugs  in 
the  gut  contents  of  a  single  fish  (Table  1).  The 
size  of  ingested  quahaugs  ranged  from  35-86 
mm  in  shell  length,  representing  quahaugs  ap- 
proximately 4-60  years  old.  Qualitative 
estimates  of  total  fresh  weights  of  quahaugs  in- 
gested per  fish  ranged  from  255.5-737.7  g;  meat 
weights  were  from  81.1-155.3  g,  respectively. 
Meat  weights  expressed  as  a  percentage  of  total 
fish  weight  (Table  1)  indicated  that  only  0.7- 
2.2%  of  fish  live  weight  is  a  food  ration  in  the 
form  of  quahaug  meat. 

It  is  realized  that  the  sample  size  is  small,  but 
the  fact  that  ocean  quahaugs  comprised  over 
85%  (by  wet  weight)  of  the  diet  of  these  woiffish 
is  of  interest.  This  is  supported  by  the  insignifi- 
cant amounts  of  macro-invertebrate  fauna  in 
the  stomaches  of  fish  with  quahaugs.  In  con- 
trast, the  sanddollar  {E.  parma)  constituted 
100%  of  the  diet  of  one  fish.  This  finding  sup- 
ported our  reconnaissance  of  the  study  site 
which  revealed  a  general  lack  of  macro- 
epibenthos.  These  results  further  suggest  a 
possible  selection  preference  for  quahaugs  by 
woiffish,  although  other  items  of  marginal  food 
value  are  also  eaten.  The  slow  growth  oi Arctica 
islandica  (Murawaski  et  al.  1982)  makes  it 
available  to  woiffish  for  50-60  years  after  the 
clam  settles  on  the  bottom.  This  is  a  long  period 
of  potential  exposure  to  predation  by  woiffish, 
l)ut  the  duration  of  the  life  span  infers  low 
natural  mortality.  This  in  part  retlects  low  levels 
of  woiffish  al)undance,  their  ability  to  locate  and 
capture  prey  as  well  as  the  clams  ability  to  avoid 


CO 

> 

o 
Z 


S 


>■, 

13 
c 


3 

c/: 


< 


^  ^ 


%-s. 


3  !i 

C   0) 


3  ^ 

O    4t 


8S 


5!_ 


x:    4)     •  — 


O     C    4J   ~- 


e  %~ 


fN   <  CO  r- 
.   \     .     . 

-N     2    •-    O 


f^    tn     r~     rf 

o  in  ■-  r-  1 
>-  in  CD  CO 


en  r-   i£)  v£i 


in   r-    rs)    •? 


o   o    *T   o 


O^    li)    •-   I*)  o 


f*l    O^    'T    CD 

r^   ^   o   vD 
CD  r-   o>  1-1 


^    O     I 


CD   Z  r-  o   Z 


o   •-   •-   o   o 


•-  (N  m  «»  in 


Ul 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    129 


predation. 

Under  the  assumption  that  a  minimum  daily 
ration  of  only  5%  body  fresh  weight  is  required 
for  maintenance  (pers.  comm.  J.  Castell,  DFO, 
Halifax,  N.S.)  the  consumption  of  quahaugs  by 
wolffish  in  this  study  was  considerably  lower 
than  this  minimum  value  and  ranged  from  0.7  to 
2.2%  (Table  1).  This  is  probably  a  reflection  of 
low  availability  of  ocean  quahaugs  in  the  study 
area.  Rowell  and  Chaisson  (1983)  estimated  that 
the  mean  density  of  ocean  quahaugs  in  the  im- 
mediate study  area  was  3.8  m^  Over  a  much 
larger  area  of  commercial  potential  in  the  same 
region  (12  km^)  they  estimated  a  mean  density 
of  5.4  m\  In  the  study  area,  the  results  indicated 
that  wolffish  probably  prey  on  quahaugs  equal 
to  the  number  in  one  m^  each  day.  Consequent- 
ly, even  in  low  abundance  wolffish  may  exert 
considerable  impact  on  the  commercial  densities 
of  quahaugs.  It's  impact  on  quahaug  beds  in 
areas  suitable  for  commercial  exploitation  may 
be  higher  but  more  research  is  needed  to  sup- 
port such  a  hypothesis. 

Acknowledgments 

We  wish  to  thank  P.  Woo,  I&MP,  DFO;  D. 
Chaisson,  Bio-Atlantech;  and  J.  Ropes,  U.  S. 
Dept.  Commer.,  NOAA,  NMFS,  Woods  Hole, 


Mass.,  for  their  assistance  in  sample  collection, 
analysis  and  manuscript  review. 

LITERATURE  CITED 

Arntz,  W.  E.  1974.  A  contribution  to  the  feeding  ecology  of 
juvenile  cod  (Gadus  morhua  L.)  in  the  western  Baltic. 
Rapp.  P.V.  Reun.  Cons.  int.  Explor.  Mer.  166:13-19. 

1978.  The  food  of  adult  cod  {Gadun  morhua  L.)  in 

the  western  Baltic.  Meeresfarsch.  26(1977-78):60-69. 

Bigelow,  H.  B.  and  W.  C.  Schroeder.  1953.  Fishes  of  the 
Gulf  of  Maine.  Fish  Bull.  U.S.  53,  577  pp. 

Bowman,  R.  E.,  R.  0.  Maurer  Jr.  and  J.  Murphy.  1976. 
Stomach  contents  of  twenty-nine  fish  species  from  the  five 
regions  in  the  Northeast  Atlantic.  Data  report  Northeast 
Fisheries  Centre,  Woods  Hole.  Mass.  Lab.  Ref.  No.  76-10, 
37  pp. 

Gill,  T.,  1911.  Notes  on  the  structure  and  habits  of  the  wolf- 
fishes.  Proceedings  U.S.  National  Museum  39(1782): 
157-187. 

Murawski,  S.  A.,  J.  W.  Ropes,  and  F.  M.  Serchuk,  1982. 
Growth  of  the  ocean  quahaug,  Arctica  islandica.  in  the 
Middle  Atlantic  Bight.  Fish  Bm.//.80:21-34. 

Ropes,  J.  W.  and  S.  A.  Murawski.  1983.  Maximum  shell 
length  and  longevity  in  ocean  quahaugs,  A rctica  islandica. 
Linne.  ICES/CM.  1983/K:32,  Shellfish  Comm.  8pp. 

Rowell,  T.  W.  and  D.  R.  Chaisson.  1983.  Distribution  and 
abundance  of  the  ocean  quahaug  {Arctica  islandica).  and 
Stimpson's  Surf  Clam  {Spisula  polymyma)  resource  on  the 
Scotian  Shelf.  Can.  Ind.  Rep.  Fish.  Aquatic  Sci.  142:v  + 
75  pp. 

Verrill,  A.  E.  1871.  On  the  food  habits  of  some  of  our  marine 
fishes.  Amer.  Nat  5:397-400. 


ATTENTION:  RESEARCHERS  AND  SMELL  COLLECTORS.   ANNOUNCING 

MALACOLOGY 
DATA       NET 

ECOSEARCH     SERIES 


A  new  and  unique  journal  specializing  in  rapid  publication 
of  worthy  papers  on  marine,  freshwater,  and  terrestrial 
moUusks.   Special  attributes  are: 

>  6  numbers  per  year  (150-175  pages); 

>  Peer  review  and  publication  within  8  weeks; 

>  Inexpensive:   $14.50/year,  $18.00  foreign; 


>    Exchange  notices  (up  to  30  words)  printed 
free,  once  a  year,  for  subscribers; 

Most  journals  take  a  year  to  publish  manuscripts. 
That  is  too  long.    By  facilitating  rapid  publication. 
Data  Net  provides  unusual  opportunities  for  timely 
interactions  between  amateurs  who  are  willing  to 
contribute  specimens  and  experience  with  professionals 
who  are  at  the  cutting  edge  of  science. 

Please  mail  in  your  order  today.    Make  your  check 
payable  to  ECOSEARCH,  Inc.   This  new  venture 
needs  your  support.   Thank  you. 

Arthur  II.  Clarke,  Editor 

(former  AMU  President 

and  Associate  Curator, 

Smithsonian  Institution) 

ECOSEARCH,  IMC. 

Etokigital  Sufveyi  ^nd  Rese<*(ch 

325  E.  Bayview,  Portland,  Tx.  78374 


130    THE  NAUTILUS  October  31,  1986  Vol.  100(4) 

FRESHWATER  MOLLUSKS  OF  THE  DOMINICAN  REPUBLICi 


Jose  D.  Gomez,  Mercedes  Vargas  and  Emile  A.  Malek 

Universidad  Autonoma  de  Department  of  Tropical  Medicine 

Santo  Domingo  Tulane  Medical  Center 

Dominican  Republic  New  Orleans,  LA  70112 

ABSTRACT 
A  survey  of  the  freshwater  mollusks  of  the  Dominican  Republic  revealed  the 
presence  of  27  species,  2U  of  them,  gastropods  and  3  bivalves.  There  were  10  species 
of  the  family  Planorbidae  and  of  these  k  were  species  o/Biomphalaria,  namely,  B. 
glabrata,  the  intermediate  host  of  Schistosoma  mansoni,  B.  havanensis,  B. 
helophila,  and  B.  straminea.  The  role  of  the  latter  3  species  in  transmission  of 
schistosomiasis  in  the  country  is  under  investigation.  Other  families  found  were 
Lymnaeidae,  Physidae,  Ampullariidae  and  Thiaridae  with  2  species  each,  and 
Ancylidae.  Hydrobiidae  and  Pleuroceridae  with  1  species  each.  There  were  3 
species  of  the  Neritidae. 


There  have  been  only  a  few  reports  on  the 
freshwater  mollusks  of  the  Dominican  Republic. 
Probably  the  first  was  by  Crosse  (1891),  who 
listed  the  land  and  freshwater  snails  of  the 
island  of  Santo  Domingo  (Hispaniola),  but  the 
nomenclature  he  used  naturally  does  not  agree 
with  that  used  today.  Other  writers  have 
reported  the  occurrence  of  the  snail  Bi.07nph.a- 
laria  glabrata  (Say,  1818)  in  either  a  few  or 
several  habitats  (Ponce  Pinedo,  1947;  Olivier  et 
al.,  1952;  Etges  and  Maldonado,  1969; 
Schneider  et  al.,  1985),  and  still  others  have 
listed  a  few  freshwater  mollusks  associated  with 
B.  glabrata  (Gomez,  1973;  Vargas  and  Gomez, 
1976;  Schneider  et  al.,  1985).  The  lymnaeid  in- 
termediate hosts  of  Fasciola  hepatica  were 
reported  upon  by  some  workers  (Alvarez  and 
Cordeiro,  1977;  Gomez  et  al.,  1986). 

In  this  paper  we  include  the  results  of  surveys 
for  the  freshwater  mollusks  of  the  country  car- 
ried out  for  the  last  five  years  by  the  Institute  of 
Research  in  Bilharzia  of  the  Autonomous  Uni- 
versity of  Santo  Domingo  and  by  the  junior 
author. 

Materials  and  Methods 

Hydrographic  maps  were  consulted  to  locate 


•Contribution  from  the  Institute  de  Investigaciones  en 
Bilharzia,  U.A.S.D.  Dominican  Republic  and  the  World 
Health  Organization  Collaborating  Center  for  Applied 
Medical  Malacology,  Tulane  Univer.sity. 


freshwater  bodies  and  detailed  sectional 
regional  maps  were  prepared  to  locate  the 
various  habitats.  Identification  of  the  specimens 
was  based  on  information  from  the  Pan 
American  Health  Organization  (PAHO)  Guide 
(1968),  Malek  (1985),  and  various  literature  on 
mollusks  from  the  Great  and  Lesser  Antilles. 

Included  in  the  information  obtained  for  each 
body  of  water  was  the  associated  fauna  and 
flora  as  well  as  the  rainfall  cycle  and  other 
physical  and  chemical  factors  influencing  the 
mollusks;  these  data  will  be  reported  in  a 
separate  paper. 

Results 

Twenty-four  species  of  freshwater  gastropods 
were  collected  during  this  survey;  these  be- 
longed to  nine  families  (Table  1).  Three  species 
of  bivalves  were  also  found.  Among  the 
gastropod  families  there  were  10  species  of 
Planorbidae,  of  which  Biomphalaria  helophila 
(Orbigny,  1835)  was  the  most  common  (31 
habitats),  B.  glabrata.  intermediate  host  of 
Srhisto.'^oma  mansoni  (26  habitats),  followed  by 
B.  hnvanensis  (Pfeiffer,  1839)  (15  habitats),  and 
B.  straminea  (Dunker,  1848)  (1  habitat).  B. 
helophila  and  B.  hnvanensis  are  potential  hosts 
for  S.  mansoni.  and  B.  straminea  is  a  known 
transmitter  of  this  schistosome  in  South 
America. 

Other  families  encountered  were  Lymnaeidae, 
Physidae,  Ampullariidae  and  Thiaridae  with  2 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    131 


TABLE  1.  Families  and  species  of  gastropods  and  their  loca- 
tions in  the  Dominican  Republic. 

Planorbidae 

Biomphala ria  glnhrnta 
Hato  Mayor;  Santo  Domingo;  La  Vega;  San  Cristobal; 
Bayaguana;  Haina;  El  Seibo;  Sabana  de  la  Mar;  Guerra; 
San  Jose  de  los  Llanos;  San  Pedro  de  Macoris;  San  Fco  de 
Macoris;  Arenoso;  Higuey;  Miches;  Nisibon;  Constanza; 
Jarabacoa;  Cotui;  San  Rafael  del  Yuma;  La  Romana; 
Ramon  Santana;  Santiago;  Quisqueya;  Nagua;  El  Valle. 

B.  havanensis 
Santo   Domingo;    Haina;    Bani;    Palenque;    S.   J.   de    la 
Maguana;  Las  Matas  de  Farfan;  La  Romana;  Villa  Vas- 
quez;  S.  P.  de  Macoris;  La  Vega;  Valverde;  Puerto  Plata; 
Esperanza;  Navarrete;  Villa  Gonzalez. 

B.  helophila 
Santo  Domingo;  Haina;  Valverde;  Las  Matas  de  Farfan;  S. 
J.  de  Maguana;  Barahona;  Cabral;  Las  Marias;  Neyba; 
Batey  4  and  5;  Guerra;  La  Vega;  Sabana  Rey;  Seibo;  Hato 
Mayor;  LA,  Yabiquin  River;  LA,  Yugery  River;  LA,  Yonu 
River;  Miches;  Sta.  Lucia  Stream,  Seibo;  Lebron  Stream. 
Seibo;  Pana  Pana  River;  S.F.M.,  Juda  River;  S.F.M.  El 
Gran  Estero;  S.F.M.  Arenoso;  Bhona.  Los  Patos;  Bhona, 
Quarantitre  River;  La  Vega.  .Juma  River;  Ducantamiento 
River;  Neyba;  La  Cabirma,  Neyba. 

B.  straminea 
Los  Llanos,  SPM. 

Helisomti  trivolvis 
Valverde  Mao  (canals  in  rice  fields). 

H.  foveale 
Santo  Domingo,  Distrito  Nacional;  concrete-lined  foun- 
tains,  Centro  de  los   Heroes;   concrete-lined   fountains. 
Natural  History  Museum;  Cabral  Pond,  Barahona. 

Drepaiiotrema  tucidum 
Haina;  S.  J.  Maguana;  Cabral;  Villa  Vazquez;  La  Vega; 
Puerto  Plata;  Mao;  San  Raf.  del  Yuma;  Guerra;  S.F.M., 
Casa  de  Altos;  S.F.M..  La  Estancia;  S.F.M,  El  Aguacate; 
S.F.M..  Arenoso. 

D.  cimex 
Constanza;  Haina;  Gabon  Pond,  Haina;  La  Vega,  El  Ran- 
chito;  La  Vega,  La  Cabulla;  La  Vega,  Sabana  Rey;  Val- 
verde, Guayacanes;  Mao;  Yerba  de  Guinea.  Mao;  Puerto 
Plata;  Villa  Vazquez;  Las  Lagimas,  Santiago;  Moca; 
Guerra,  D.N.;  S.F.M.,  Arenoso;  D.N.,  Botanical  Gardens. 

D.  anatinurn 
D.N.,  Botanical  Gardens;  Haina;  Km.  9  Aut,  Duarte  D.N.; 
Villa  Mella;  Mao;  S.J.  de  la  Maguana;  Cabral;  Villa 
Vazquez;  Gabon  Pond,  Haina;  Ranchito,  L.V,;  Tabagiia; 
Puerto  Plata;  Dajabon;  Moca;  San  R.  del  Yuma;  Guerra; 
Nisao;  S.F.M.,  casa  de  Alto;  San  Fco.  de  Macoris;  S.F.M. 
Arenoso;  S.F.M.,  El  Gran  Estero;  Bonao;  San  Pedro  de 
Macoris. 

D.  aeruginotiiim 
La  Vega,  El  Ranchito. 

Physidae 

Physa  cubensis 
Constanza;    La   Vega,    Gima;    La    Vega,    Sabana    Rey; 
Guayacanes,  Mao;  Tabagua;  Cerro  Gordo,  Mao;  Los  Pinos, 
Mao;  Haina. 

P.  morrnorata 
D.N.,  Botanical  Gardens;  Haina;  Hda.  La  El.strclla;  Bani; 


Sabana  Grande  de  Palen(|ue;  Guayacanes.  Mao;  Las  Matas 
de  Farfan;  Neyba,  Batey  4  and  .5;  Biran  Stream,  Barahona; 
Azua;  Pedregal,  Bani;  Jarabacoa;  Guayubin;  Villa  Vaz- 
quez; La  Vega;  Mao;  Tabagua;  Puerto  Plata;  Moca  River; 
Juan  Lopito,  Moca;  Hato  Mayor;  Martin  Avila,  LA; 
Anguilla  River,  El  Seibo;  Yonu  River,  LA;  Miches;  Santa 
Lucia  Stream;  El  Seibo;  Lebron  Stream,  Seibo;  Guerra; 
Don  Gregorio  Pond,  Nisao;  Nisao;  San  Fco.  Macoris,  Casa 
de  Alto;  SFM,  La  Estancia;  SFM,  El  Aguacate;  SFM, 
Arenoso;  Juma;  Bonao. 

Lymnaeidae 

Fossaria  cubensis 
D.N..  Botanical  Gardens;  Constanza;  Haina;  KM.  9  Duarte 
Highway,  D.N.;  Sabana  Grande  de  Palenque;  Valverde; 
Santiago;  San  J.  de  la  Maguana;  Barahona;  Las  Matas  de 
Farfan;  Azua;  Bani;  Jarabacoa;  La  Vega;  Esperanza,  Mao; 
Villa  Vazquez;  Hato  Mayor;  Miches;  Seibo;  San  Fco  de 
Macoris;  Bonao. 

Pseudosuccinea  coin  mella 
Quita  Sueno  Pond,  Haina;  D.N..  Botanical  Gardens. 

Ancylidae 

Ferrissia  irrorata 
Mao  River,  Mao;  D.N.,  Botanical  Gardens;  Cabral;  Seibo; 
San  Cristobal;  San  Juan  de  la  Maguana. 

Thiaridae 

Thiara  granifera  and  T.  tuberculntn 
D.N.,  Botanical  Gardens;  La  Vega;  Santiago;  San  J.  de  La 
Maguana;  Paraiso,  Bhona;  Cabral;  Descubierta;  Las 
Barias;  Jimani;  Neyba;  Pedernales;  Haina;  Bani;  Azua; 
Jarabacoa;  Guayubin;  Villa  Vasquez;  Sabana  de  la  Mar; 
Miches;  Nisibon;  Seibo;  Higuey;  La  Vega;  San  Pedro  de 
Macoris;  San  Fco.  de  Macoris;  Guerra;  Hato  Mayor;  Moca; 
Dajabon;  Ramon  Santana;  Puerto  Plata;  Nagua;  Castillo: 
Mao;  Samana;  Sanchez;  Bayaguana;  Nigua. 

Ampullariidae 

Marisa  comuarietis 
D.N.,  Botanical  Gardens;  Guerra;  Bayaguana;  San  Pedro 
de  Macoris;   La  Vega;  Jarabacoa;  Nigua;   Hato  Mayor; 
Nisibon;  Valverde;  Nagua. 

Ampullaria  glnuca 
Los  Llanos;  D.N.,  Botanical  Gardens;  Hato  Mayor. 

Hydrobiidae 

Pyrgophorus  parvulus 
D.N.,  Botanical  Gardens. 

Pleuroceridae 

Goniobasis  sp. 
Nigua;  D.N..  Botanical  Gardens;  Nisibon. 

Neritidae 

Neritina  sp.;  N.  pmictulata:  N.  virgiriea 
Puerto  Plata;  Haina;  Santo  Domingo;  Nisibon. 


species  each,  and  Ancylidae,  Hydrobiidae,  and 
Pleuroceridae  with  1  species  each.  Three  species 
belonging  to  the  Neritidae  were  encountered, 
and  3  species  of  bivalves:  Pisidium  puncti- 
forum.  Eupera  cubensis.  and  Anodonfa  sp.  The 
first  2  species  are  sphaeriids,  family  Sphaeri- 
idae,  and  the  third  belongs  to  the  family 
Unionidae. 


132    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


More  than  one  mollusk  shared  the  habitat  with 
at  least  one  other  moliusi<.  Thiara  spp., 
however,  did  not  occur  together  with  B. 
glabrata,  but  Marisa  cornuarietis  (Linnaeus, 
1758)  did  inhabit  the  same  waterbody  as  B. 
glabrata.  The  flora  and  fauna  in  each  habitat  did 
not  limit  the  distribution  of  mollusks.  On  the 
other  hand,  aquatic  vegetation  favored  the  oc- 
currence of  most  of  the  species.  However, 
Thiara  spp.  always  favored  habitats  with  little 
or  no  vegetation. 

Discussion 

Until  1972  B.  glabrata  occurred  in  a  total  of  1 1 
habitats  in  the  country  (Vargas  and  Gomez, 
1976),  whereas  we  found  it  in  26  habitats  in  the 
present  survey,  indicating  an  apparent  continu- 
ing dissemination.  In  addition  to  natural  factors 
which  favor  spreading,  the  movement  and  intro- 
duction of  ornamental  plants  in  parks  and  other 
locations  have  apparently  contributed  to  the 
dispersal  of  this  medically  important  species.  B. 
haimnensis  and  B.  helophila  also  show  wide 
distribution,  possibly  because  of  their  ability  to 
adapt  to  habitats  in  the  dry  region  of  the  coun- 
try. B.  glabrata  has  not  yet  invaded  areas  in  the 
south.  B.  havanensis  and  B.  helophila  are  poten- 
tial hosts  of  5.  mansoni,  and  specimens  of  B. 
havanensis  from  Haiti  have  been  infected  ex- 
perimentally (Michelson,  1976).  Specimens  of  B. 
helophila  from  Puerto  Rico  have  also  been  in- 
fected experimentally  (Richards,  1961,  1963). 

While  this  paper  was  in  preparation  a  study  by 
Sodeman  et  al.  (1985)  was  published  in  which 
they  collected  6  specimens  only  of  B.  straminea. 
in  the  river  Iguamo.  We  have  had  the  opportuni- 
ty to  collect  close  to  1000  specimens  of  B. 
straminea  from  Los  Llanos,  a  different  locality 
but  in  the  same  general  area  of  San  Pedro  de 
Macoris.  Our  specimens  showed  the  characteris- 
tic features  of  this  species  (PAHO,  1968;  Malek, 
1985),  that  is,  those  of  the  shell,  of  the  penial 
complex,  and  corrugations  on  the  vaginal  sur- 
face, but  these  corrugations  were  not  distinct  in 
all  specimens  dissected.  The  previous  known 
geographical  range  of  B.  straminea  was  Mar- 
tinique, Costa  Rica,  and  South  America  (Brazil, 
Venezuela,  Guiana).  We  are  at  present  testing 
the  susceptibility  of  our  Dominican  specimens  to 
infection  with  S.  mansoni. 

It  seems  that  Helisoma  trivolvis  (Say,  1817) 


was  introduced  and  became  established  in  the 
northeast  section  of  the  country,  and  now  oc- 
curs in  large  numbers.  However,  Helisoma 
foveale  (Menke,  1830)  is  mainly  encountered  in 
small  artificial  habitats,  such  as  small  pools  and 
fountains,  indicating  its  possible  introduction 
with  tropical  ornamental  fish.  B.  glabrata  now 
does  not  occur  in  the  northeast,  and  competition 
may  have  occurred  between  Helisoma  trivolvis 
and  B.  glabrata,  as  was  demonstrated  in  the 
laboratory  (Malek  and  Malek,  1978). 

There  were  4  species  of  the  genus  Drepano- 
trema;  the  first  was  D.  lucidum  (Pfeiffer,  1839) 
and  we  believe  that  it  is  not  D.  hoffmani  F.  C. 
Baker,  1940.  This  is  because,  first,  our 
specimens  were  never  more  than  7.5  mm  in 
diameter,  and  second,  it  seems  that/),  hoffmani 
was  described  by  F.  C.  Baker  based  on  large 
specimens  of  D.  lucidum. 

The  lymnaeid  Fossaria  cuhensis  (Pfeiffer, 
1939)  has  been  known  to  occur  on  the  island 
since  the  time  of  Crosse,  and  is  a  transmitter  of 
the  liver  fluke,  Fasciola  hepatica.  Another  lym- 
naeid, Pseudosuccinea  columella  (Say,  1817), 
was  recently  reported  to  occur  in  the  Dominican 
Republic  (Gomez  et  al.,  1986).  This  latter  species 
is  an  experimental  host  for  i^.  hepatica  in  Puerto 
Rico  (Leon-Dancel,  1970),  and  is  a  natural  host 
in  Brazil  (Ueta,  1980). 

Thiara  granifera  (Lamarck,  1822)  and  T. 
tuberculata  (Muller,  1774)  have  in  recent  years 
been  introduced  into  the  Dominican  Republic  as 
well  as  other  Caribbean  islands.  Their  popula- 
tions share  the  same  habitats  and  they  can  be 
confused  morphologically,  and  thus  are  often 
reported  in  the  Caribbean  literature  as  one 
species,  T.  granifera.  We  have  examined 
specimens  from  Puerto  Rico,  Guadeloupe,  and 
Martinique  and  they  all  represent  mixed  popula- 
tions but  predominantly  T.  tuberculata.  The 
junior  author  expressed  these  views  in  a  short 
note  (McCuUough  and  Malek,  1984).  In  a  recent 
publication  (Starmuhlner,  1985),  Thiara  tuber- 
culata was  reported  from  Guadeloupe,  Dominica 
and  Martinique.  Starmuhlner  made  an  error  in 
two  of  his  figures;  plate  8,  Figures  6,  8  and  9  are 
T.  tuberculata,  but  Figure  7  and  especially 
Figure  10  are  T.  granifera.  The  two  species  can 
be  differentiated  as  follows:  in  T.  granifera  the 
shell  is  characterized  by  distinct  and  raised  axial 
ribs  reticulating  with  spiral  threads.  Whorls  of 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    133 


the  spire  are  rather  flat-sided;  body  whorl  is 
wide  and  its  height  is  more  than  half  of  total 
height  of  shell.  In  T.  tuberculata  the  shell  usual- 
ly has  low  axial  ribs  and  distinct  and  raised 
spiral  cords  especially  on  body  whorl;  sometimes 
whorls  are  nearly  smooth  with  only  incised 
spiral  threads.  Whorls  are  evenly  rounded;  body 
whorl  is  less  in  width  than  in  T.  granifera  and  its 
height  is  less  than  half  of  shell  height;  the  shell  is 
more  slender  than  that  of  T.  granifera.  and  is 
uniformly  turreted.  The  position  of  the  brood 
pouch  is  similar  in  both  species,  as  well  as  the 
mantle  edge  bearing  finger-shaped  fringes  in 
both.  Alvarez  and  Mena  Sanchez  (1973)  re- 
ported that  T.  granifera  had  been  introduced  in- 
to the  eastern  part  of  the  Dominican  Republic 
and  at  present  it  has  spread  to  other  parts  of  the 
country  (Vargas  et  al.,  1982).  There  may  now  be 
biological  control  of  B.  glabrata  by  competition 
with  T.  tuberculata  and  T.  granifera.  Some 
habitats  previously  occupied  by  B.  glabrata  now 
harbor  only  the  two  melaniid  species.  According 
to  Ferguson  (1978),  T.  granifera  competes  suc- 
cessfully with  B.  glabrata  in  Puerto  Rico.  In  St. 
Lucia,  four  field  trials  demonstrated  that  B. 
glabr-ata  was  eliminated  from  marshes  and 
streams  6-22  months  after  the  introduction  of 
T.  granifera  (Prentice,  1983). 

Marisa  corriuarietis.  an  operculate  ampul- 
lariid,  was  introduced  into  the  Dominican 
Republic  by  the  personnel  of  the  Health  Depart- 
ment in  charge  of  the  control  of  schistosomiasis 
as  a  competitor  of  B.  glabrata.  because  of  the 
encouraging  result  in  nearby  Puerto  Rico  (see 
review  by  Ferguson,  1978).  In  our  experience 
the  two  snails  now  occur  together  in  some  habi- 
tats in  the  Dominican  Republic,  and  if  M.  cor- 
nuarietis  did  succeed  in  biological  control  in 
Puerto  Rico  this  has  taken  place  only  in  certain 
situations,  for  example,  small  ponds  and  reser- 
voirs. 

The  bivalves  of  the  Dominican  Republic  have 
not  received  much  attention.  Anodonta  sp. 
(family  Unionidae)  was  recently  introduced  into 
the  country  and  is  now  present  in  the  fish 
{Tilapia  and  carp)  ponds  of  the  Ministry  of 
Agriculture  in  Nigua,  close  to  the  capital,  Santo 
Domingo. 

The  main  species  of  freshwater  mollusks  of 
the  Dominican  Republic  are  also  present  on  cer- 
tain other  Antillean  islands.   In  Haiti,   which 


shares  the  island  of  Hispaniola  with  the 
Dominican  Republic,  Robart  et  al.  (1976) 
reported  16  species;  in  Puerto  Rico,  van  der 
Schalie  (1948)  reported  14  species;  In 
Guadeloupe,  Pointier  (1974)  listed  21  species, 
and  in  Martinique,  Guyard  and  Pointier  (1979) 
listed  19  species.  In  this  paper  we  report  27 
species  to  be  present  in  the  Dominican  Republic, 
but  some  of  these  are  of  recent  introduction. 

LITERATURE  CITED 

Alvarez,  J.  M.  V.,  and  Cordero,  E.  T.  C.  1977.  Fascioliasis 
bovina  en  la  Republica  Dominicana.  Publ.  Universidad 
Autonoma  S.  Domingo.  Vol.  224.  Col.  Ciencia  y  Tech.  No. 
5:1-189. 

Alvarez,  J.  M.  V.,  and  Mena  Sanchez,  V.  1973.  Hallazgo  en 
Dajabon,  Republica  Dominicana  del  caracol  de  agua  dulce 
"Thiara"  spp.  de  importancia  medica.  Revista  de  la 
Faciiltad  de  Ciencias  Agroitomicat:  y  Veterinarias 
2:.53-.54. 

Crosse,  H.  1891.  Faune  malacologique  terrestre  et  fluviatile 
de  rile  de  Saint  Domingue.  Jour.  Conrhyliol.  39:69-211. 

Etges,  F.  J.,  and  Maldonado,  J.  R.  1969.  The  present  status 
of  bilharziasis  in  the  Dominican  Republic.  Malacologia 
9:40-41. 

Ferguson,  F.  F.  1978.  The  role  of  biological  agents  in  the 
control  of  schistosome-bearing  animals.  Atlanta,  Georgia, 
U.S.A.,  U.S.  Department  of  Health,  Education  and 
Welfare,  Center  for  Disease  Control,  107  pp. 

Gomez,  J.  1983.  Distribucion  geografica  del  Biomphalaria 
glabrata  (huesped  intermediario  del  Schistosoma  mansoni) 
en  Republica  Dominicana.  Thesis  for  Licenciado  en 
Biologia,  Universidad  Autonoma  de  Santo  Domingo. 

Gomez,  J.,  Vargas,  M.  and  Malek,  E.  A.  1986.  P-'seudosiic- 
cinea  columella  (Lymnaeidae)  in  the  Dominican  Republic 
and  transmission  of  fascioliasis  in  the  Caribbean  region. 
The  Nautilus  100:66-68. 

Guyard,  A.  and  Pointier,  J.  P.  1979.  Faune  malacologique 
dulcaquicole  et  vecteurs  de  la  schistosome  intestinale  en 
Martinique  (Antilles  Francaises).  Ann.  Parasitol.  (Paris) 
54:193-205. 

Leon-Dancel,  D.  1970.  Life  history  of  Lymnaea  columella 
(Say)  and  its  e.xperimental  infection  with  Fasciola  hepatica 
(L.).  Jour.  Agric.  Umv.  Puerto  Rico.  54:297-305. 

Malek,  E.  A.  1985.  Snail  hosts  of  Schistosomiasis  and  Other 
Snail-transmitted  Diseases  in  Tropical  America:  A 
Manual.  Pan  American  Health  Organization,  Washington, 
D.  C,  Scientific  Publication  No.  478,  1-325. 

Malek,  E.  A.,  and  Malek,  R.  R.  1978.  Potential  biological 
control  of  schistosomiasis  intermediate  hosts  by  helisome 
snails.  The  Nautilus  92:15-18. 

McCullough,  F.  S.,  and  Malek,  E.  A.  1984.  Notes  on  the  mol- 
luscan  intermediate  hosts  oi  Paragonimus  spp.  and  their 
possible  role  in  biological  control.  Ann.  Trap.  Med.  Para- 
sitol. 78:339-340. 

Michelson,  E.  H.  1976.  A  potential  intermediate  host  of 
Schistosoma  mansoni  from  Haiti.  Jour.  Parasitol. 
62:648-649. 

Olivier,  L.,  Vaughn,  C.  M.  and  Hendricks,  J.  B.  1952.  Schis- 


134    THE  NAUTILUS 


October  31,  1986 


Vol.  100  (4) 


tosomiasis  in  an  endemic  area  in  the  Dominican  Republic. 
ArMT.  Jour.  Trop.  Med.  Hyg.  1:680-687. 

Pan  American  Health  Organization.  1968.  A  Guide  for  the 
Identification  of  the  Snail  Intermediate  Hosts  of  Schisto- 
somiasis in  the  Americas.  PAHO  Scientific  Publication  No. 
168,  1-122. 

Pointier,  J.  P.  1974.  Faune  malacologique  dulcaquicole  de 
rile  de  la  Guadeloupe  (Antilles  Francaises).  Bull.  Mus. 
Hisl.  Natur..  Paris,  3rd  Ser.,  No.  235,  Zool.  159:905-933. 

Ponce  Pinedo,  A.  M.  1947.  Schistosomiasis  mansoni  in  the 
Republic  of  Santo  Domingo  with  a  report  of  six  cases 
studied.  Puerto  Rico  Jour.  Publ.  Hlth.  Trap.  Med. 
22:308-324. 

Prentice,  M.  A.  1983.  Displacement  of  Biomphalarin  gin- 
brata  by  the  snail  Thiarn  granifera  in  field  habitats  in  St. 
Lucia,  West  Indies.  .Ann.  Trop.  Med.  Parasitol.  77:51-59. 

Richards,  C.  S.  1961.  Another  potential  intermediate  host 
snail  for  Schistosoma  mansoni  in  Puerto  Rico.  Jour. 
Parasitol.  47:64. 

1963.  Infectivity  of  Schistosoma  mansoni  for 

Puerto  Rican  mollusks,  including  a  new  potential  inter- 
mediate host.  Ainer.  Jour.  Trop.  Med.  Hyg.  12:26-33. 

Robart,  G.,  Mandahl-Barth,  G.  and  Ripert,  C.  1976.  Inven- 
taire,  repartition  geographique  et  ecologie  des  mollusques 
dulcaquicoles  d'Haiti  (Caraibes).  Hnliotis.  8:159-171. 


Schneider,  C.  R.,  Hiatt,  R.  A.,  Malek,  E.  A.  and  Ruiz-Tiben, 
E.  1985.  Assessment  of  schistosomiasis  in  the  Dominican 
Republic.  Public  Hlth.  Rejjorts  100:524-530. 

Sodeman,  W.  A.,  Rodrick,  G.  E.,  Paraense,  W.  L.  and 
Vargas  de  Gomez,  M.  1985.  Biomphalaria  straminea  and 
other  planorbids  in  the  Dominican  Republic.  Mem.  Inst. 
Oswaldo  Cruz,  Rio  de  Janeiro  80:453-456. 

Starmuhlner,  F.  1985.  Erstfunde  von  drei  Arten  von  Suss- 
wasser  Gastropoden  auf  den  Inseln  Guadeloupe,  Dominica 
und  Martinique  (Kleine  Antiilen).  Heldin  1:55-58. 

Ueta,  M.  T.  1980.  Ocorrencia  de  infeccao  natural  Ae Fasciola 
hepatica  Linnaeus,  1758  em  Lymnaea  columella  Say,  1817 
no  Vale  de  Paraiba,  SP,  Brasil.  Re^i.  Saude  Publica 
14:230-233. 

Van  der  Schalie,  H.  1948.  The  land  and  fresh  water  mollusks 
of  Puerto  Rico.  Misc.  Publ.  Mus.  Zool.  Univ.  Michigan, 
No.  70. 

Vargas  Castro,  M.,  and  Gomez,  J.  1978.  Algunos  aspectos  de 
la  distribueion  del  Biomphalaria  glabrata  en  la  Republica 
Dominicanas.  Ciencia,  Organo  de  la  Direccion  de  Investi- 
gaciones  Cientificas  de  la  U.A.S.D.,  3:87-94. 

Vargas,  M.,  Gomez,  J.  D.  and  Guerrera,  N.  1982.  Distribu- 
eion en  la  isla  de  Santo  Domingo  de  Thiara  granifera 
(Lamarck),  Mollusca:  Melanoide.  Rev.  Dam.  Microbiol. 
Parasitol.,  1:14-18. 


A  PREHISTORIC  ABORIGINAL  FRESHWATER  MUSSEL  ASSEMBLAGE 
FROM  THE  DUCK  RIVER  IN  MIDDLE  TENNESSEE 

Paul  W.  Parmalee  and  Walter  E.  Klippel 

Department  of  Anthropology 

University  of  Tennessee 

Knoxville,  TN  37996 

ABSTRACT 

A  tofiil  of  761  valves  of  freshwater  mussels,  representing  a  minimum  of  28 
species,  was  identified  from  a  prehistoric  ahoriginal  cave  site  located  along  the 
Duck  River,  Maury  County,  Tennessee.  Two  thirds  of  the  mussels  are  Cumber- 
landian  forms  and.  as  a  group  and  based  on  known  habitat  requirements,  suggest 
that  this  stretch  of  the  Duck  River  for  some  period  between  ca.  7.000  and  1,000  BP 
was  more  shallow  than  at  present  with  numerous  riffles  and  swift  current. 
Recovery  of  numerous  specimens  of  Pegias  fabula  from  aboriginal  sites  located 
along  a  ca.  150  km  stretch  of  the  Duck  River  irfects  an  extensive  prehistoric 
population  of  this  mussel  in  Middle  Tennessee.  This  prehistoric  assemblage  is 
discussed  in  relation  to  Duck  River  m.ussel  faunas  reported  over  the  last  60  years. 


Several  small  rockshelters  situated  in  the 
Duck  River  bluffs  were  tested  for  aboriginal 
occupation  during  archaeological  survey  work  in 
1978  in  the  proposed  Tennessee  Valley  Author- 
ity  Columbia   Reservoir  area.   One   of  these, 


Cheek  Bend  Cave  (40MU261),  is  located  approx- 
imately 13  km  ESE  of  the  city  of  Columbia, 
Maury  County  (Fig.  1),  and  testing  (three  1  x  2  m 
excavation  units)  showed  it  to  contain  stratified 
deposits  of  bone  and  shell  to  a  depth  of  approxi- 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    135 


TENNESSEE 


10  20  JOk 


FIG.  1.  The  Duck  River  in  Middle  Tennessee  with  the  loca- 
tion of  Cheek  Bend  Cave  and  other  aboriginal  sites  contain- 
ing naiads  discussed  here. 


mately  4.5  m.  On  the  basis  of  the  vertebrate 
species  composition  within  and  among  strata, 
differing  fill  zones,  and  certain  other 
stratigraphic  features,  the  deposit  reflects  two 
distinct  and  well-defined  major  episodes  of  fill. 
The  top  2  m  represent  the  Holocene  stage  and 
contain  remains  of  extant  modern  species.  In 
contrast,  the  faunal  assemblage  in  the  bottom 
2  m,  although  it  includes  some  species  that  still 
inhabit  the  cave  area,  contains  many  such  as  the 
prairie  chicken,  13-lined  ground  squirrel  and 
pocket  gopher  that  are  now  extirpated  and 
reflect  a  prairie  habitat  (Parmalee  and  Klippel, 
1981)  and/or  a  boreal  environment,  species  such 
as  red-backed  and  yellow-cheeked  voles,  red 
squirrel,  northern  flying  squirrel,  and  arctic  and 
water  shrews  (Klippel  and  Parmalee,  1982). 

Matrix  from  two  additional  one-meter-square 
columns  was  removed  during  1982  and  1983. 
Only  the  vertebrate  remains  from  the  Pleisto- 
cene strata  have  undergone  preliminary  iden- 
tification, but  all  freshwater  mussel  valves  from 
the  Holocene  strata  were  removed  and  incor- 
porated with  the  1978-1979  specimens  for  this 
study.  Although  terrestrial  gastropods  occurred 
in  all  strata,  aquatic  gastropods,  freshwater 
mussel  shells  and  prehistoric  human  artifacts 


were  encountered  only  in  the  Holocene  strata. 
Mussels  were  commonly  used  as  a  food  resource 
by  prehistoric  Indians  in  eastern  North  America 
(Parmalee  and  Klippel,  1974)  and  in  all  proba- 
bility most  if  not  all  of  the  naiads  present  in  this 
cave  deposit  were  gathered  by  aboriginal  groups 
occupying  the  site. 

The  majority  of  valves  were  well-preserved 
with  some  in  the  top  two  strata  still  retaining 
remnants  of  the  periostracum,  but  also  a  large 
number  of  shells  were  broken  with  only  portions 
of  the  hinge  line  remaining.  Because  of  this  lat- 
ter condition,  an  additional  555  valve  fragments 
could  not  be  identified  (Table  1).  All  species 
determinations  were  made  using  comparative 
reference  specimens  in  the  collections  of  the 
Section  of  Zooarchaeology,  Department  of 
Anthropology,  University  of  Tennessee,  Knox- 
ville;  all  mollusk  and  vertebrate  remains 
recovered  from  Cheek  Bend  Cave  are  housed  in 
the  Department  of  Anthropology. 

The  Duck  River  and  Molluscan  Studies 

Interest  by  American  malacologists  in  the 
varied  and  abundant  freshwater  molluscan 
fauna  once  present  in  most  major  river  systems 
in  eastern  North  America  began  well  over  a  cen- 
tury ago.  These  beginning  efforts  centered 
around  taxonomy,  and  emphasis  was  placed  on 
describing  and  naming  species  new  to  science. 
By  the  1850s  the  majority  of  "new"  species  had 
been  described  and  the  tendency  then  was  to 
treat  the  total  molluscan  fauna  of  a  specific 
region  or  river  system.  List  of  Shells  Collected  in 
Central  Tennessee  by  Hinkley  and  Marsh  (1885) 
is  only  one  of  many  such  references,  but  we  cite 
it  because  it  provides  not  only  a  list  of  species 
known  from  the  Duck  River  (".  .  .  the  Duck  River 
and  adjacent  country  at  Columbia,  Maury  Co.") 
at  that  time  but  also  makes  reference  to  the 
abundance  of  many  species  and  the  habitat  in 
which  they  occurred.  In  addition,  the  following 
description  by  Hinkley  and  Marsh  (1885:2) 
seems  appropriate:  "The  Duck  River,  at  Colum- 
bia, is  a  pretty  and  rather  picturesque  stream, 
the  limestone  bluffs  along  its  banks  having  been 
sculptured  by  the  action  of  rain  and  frost  into 
various  shapes,  in  places  overhanging  the  water 
and  forming  quite  a  cavern  underneath.  The 
water  is  shallow,  swift,  and  clear.  .  ."  See  Fig.  2. 

After  collecting  the  Duck  River  in  1921,  1922, 


136    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


FIG.  2.  View  of  the  Duck  River,  Maury  County,  winter 
1978-79,  approximately  100  m  upstream  from  Cheek  Bend 
Cave. 

and  1923,  Ortmann  (1924)  published  a  compre- 
hensive treatment  of  the  naiad  fauna  that  in- 
cluded 63  species  and  forms  which  he  confirmed 
and  7  that  "should  be  credited  to  the  Duck 
River"  based  on  previous  published  reports. 
More  recent  studies  by  Isom  and  Yokley  (1965) 
and  van  der  Schalie  (1973)  have  pointed  up  the 
fact  that  the  Duck  River  naiad  assemblage  has 
now  been  reduced  by  nearly  one-fourth  to  about 
45  species  or  forms.  Completion  of  the  Columbia 
dam  and  reservoir  will  bring  about  the  extirpa- 
tion of  most  species  in  the  effected  stretches  of 
the  river  and  will  probably  eliminate  two 
species,  now  known  to  inhabit  only  this  locale, 
from  the  entire  river  system.  Ahlstedt  (1981),  in 
a  recent  survey  of  the  molluscan  fauna  of  the 
Duck  River  between  the  Normandy  and  Colum- 
bia dams,  found  a  drastic  decline  in  the  numbers 
and  abundance  of  species  compared  with  earlier 
surveys  in  1965  and  1973.  The  section  of  the 
Duck  River  above  Columbia  contains  the  last 
known  populations  in  that  river  of  such  species 
as  Quadrula  intermedia  and  Lemiox  rimosus.  In 
1982  TVA  biologists  transplanted  ca.  4,000  in- 
dividuals of  L.  rimosus  into  three  other  rivers 
(and  at  one  location  in  the  upper  Duck  River 
above  the  Normandy  reservoir)  in  hopes  of  re- 
establishing this  endangered  species  in  known 
habitats  where  it  once  occurred.  Success  or 
failure  of  this  ongoing  program  has  not  yet  been 
fully  evaluated  (S.A.  Ahlstedt,  pers.  commun.). 

Cheek  Bend  Cave  Mussel  Assemblage 

Although  the  shell  was,  for  the  most  part,  ex- 


ceptionally well  preserved,  the  disintegration  of 
the  periostracum  in  the  majority  of  specimens 
and  consequently  loss  of  all  color  (including  the 
nacre  in  most)  and  pattern  made  specific  iden- 
tification of  valves  representing  certain  groups 
(e.g.  Villosa.  Epioblasrna)  impossible  or  ques- 
tionable at  best.  For  example,  Ortmann  (1924: 
52)  commented  that  "The  males  of  D. 
f  =  Dysyioyriia  =  Epioblasm.aJ  florentina  walkeri, 
closely  resemble  those  of  the  next  species  {cap- 
saeformis),  but  they  differ  chiefly  in  color." 
Shells  of  species  as  distinct  as  Lampsilis  ovata 
and  Actinonaias  pectorosa,  when  fresh,  may  be 
impossible  to  distinguish  when  only  the  dorsal 
portion  of  the  valve  (hinge  line  with  lateral  and 
pseudocardinal  teeth)  is  preserved -as  in 
archaeological  or  cave  contexts. 

Nevertheless,  those  specimens  that  could  be 
identified  provide  an  interesting  record  of  the 
naiad  species  that  inhabited  the  Duck  River  in 
Middle  Tennessee  in  prehistoric  times  (on  which 
there  is  no  published  data)  and  which,  in  turn, 
reflect  river  conditions  at  the  time  they  were 
collected.  Valves  of  Epioblasrna  capsa^formis 
(some  of  which  may  be  E.  florentina)  comprised 
nearly  20%  of  the  identified  naiads  recovered  in 
the  excavation  units;  at  the  time  Ortmann  col- 
lected the  Duck  River  at  Columbia/Leftwich/ 
Lillard  Mill/Willhoite  it  was  "Rather  abundant- 
New  [previously  unreported]  for  Duck  River" 
(Ortmann  1924:53).  Although  still  found  in  the 
Duck  River  above  Columbia,  it  is  uncommon  and 
local  in  occurrence.  We  collected  several  dead 
mature  specimens  below  Lillard  Mill  that  were, 
unlike  those  from  the  cave,  extremely  large  and 
thick-shelled.  The  spike,  Elliptio  dilatata,  most 
of  which  were  small,  thin-shelled  (for  the 
species)  and  compressed -suggestive  of  a 
shallow  small  stream  habitat  -  comprised  about 
12%  of  the  cave  naiad  assemblage. 

Ptychobranchus  subtentum  is  another  species 
that  was  apparently  common  in  the  Duck  River 
near  the  cave  site  in  prehistoric  times;  107 
valves  (14%)  were  identified  from  the  deposit.  It 
was  evidently  uncommon  at  Columbia/Leftwich/ 
Normandy  when  Ortmann  (1924:40)  made  his 
1921-1923  collections,  reporting  "Only  a  few 
specimens  found."  His  further  comment  that  it 
is  "Most  abundant  in  smaller  streams  above  the 
range  of  Ptychobranchui^  fasciolare,  and  this 
seems  to  hold  good  also  in  Duck  River"  is  of  par- 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    137 


ticular  interest.  This  observation,  coupled  with 
the  numerous  valves  of  P.  subtentum.  and  other 
species  (e.g.  Medionidus  conradicus)  or  forms 
from  the  cave  that  are  indicative  of  a  head- 
waters and/or  small  stream  habitat,  suggests 
that  that  section  of  the  Duck  River  flowing  in 
close  proximity  to  Cheek  Bend  Cave  during  the 
mid-Holocene  was  smaller  and  more  shallow 
than  it  is  today. 

Another  species  represented  in  the  cave 
deposit  (Stratum  VII),  namely  Pegias  fabula,  is 
also  indicative  of  a  fast-flowing,  shallow,  small 
stream  habitat.  Archaeological  surface  collec- 
tions were  obtained  from  a  Middle  to  Late 
Archaic  (ca.  7,000-4,000  BP)  shell  midden  at  the 
Ervin  Site  (40MU147)  during  1979.  This  pre- 
historic site,  situated  ca.  29  km  upstream  from 
Cheek  Bend  Cave  in  Maury  County  (Fig.  1),  pro- 
duced 15  specimens  of  P.  fabula  out  of  1339 
identified  valves  (Hofman,  nd).  During  the 
1982-1984  excavations  at  the  Hays  site 
(40ML139),  ca  40  km  upstream  from  Cheek 
Bend  Cave  in  Marshall  County,  huge  quantities 
of  aquatic  gastropods  and  freshwater  mussel 
valves  were  removed  from  this  Middle  to  Late 
Archaic  (ca.  7,000-4,000  BP)  shell  midden.  In- 
cluded among  the  valves  of  the  32  species  iden- 
tified from  this  site  thus  far  (Turner,  nd)  were 
104  specimens  of  P.  fabula.  Even  farther 
upstream,  ca.  134  km  above  Cheek  Bend  Cave, 
Robison  (nd)  identified  43  valves  of  P.  fabula 
(from  a  sample  of  2,545  shells)  from  the  Shofner 
site  (40BD55),  Bedford  County,  a  Middle 
Woodland  village  dating  ca.  AD  600-500. 
Archaeological  sites  on  tributaries  of  the  Duck 
River  have  also  produced  P.  fabula;  one 
specimen  was  recovered  from  Woodland  and 
one  from  Middle  Archaic  strata  (266  valves  iden- 
tified) at  the  McCollum  Rockshelter  (40MU390) 
along  Fountain  Creek  in  Maury  County.  Two 
additional  specimens  (135  valves  identified) 
were  recovered  from  Goatcliff  Rockshelter 
(40MU436)  along  the  same  Duck  River  tributary 
in  what  appears  to  be  Woodland  context 
(O'Hare,  nd).  Excavation  of  these  rockshelters 
took  place  during  1980-1981  .  A  single 
specimen  of  P.  fabula  in  the  United  States  Na- 
tional Museum  collections  ("Duck  River,"  USNM 
Lot  No.  86229,  catalogued  Feb.  1888:  Paul 
Greenhall,  pers.  comm.,  April  1986)  appears  to 
be  the  only  documented  historic  record  of  this 


naiad  for  the  Duck  River  (Clarke,  1981). 

In  a  recent  paper  Starnes  and  Starnes  (1980) 
report  a  viable  population  of  P.  fabula  from  the 
Little  South  Fork  Cumberland  River,  Kentucky, 
a  small  fast-flowing  stream  ca.  20-25  m  wide 
with  an  average  water  depth  of  about  20  cm  at 
low  river  stages.  Of  additional  interest  is  the 
fact  that  the  associated  mussel  assemblage  in- 
cludes P.  subtenturn,  P.  fasciolare,  E.  dilatata, 
M.  conradicus.  Villosa  iris,  Villosa  taeniata, 
Villosa  vanuxemensis  and  Lampsilis  fasciola. 
Although  valves  of  Alasmidonta  viridis,  Toxo- 
lasyna  cylindrellus  and  Lemiox  rimosus  were 
not  numerous  in  the  Cheek  Bend  Cave  deposit, 
the  presence  of  these  species  is  also  indicative  of 
a  small  stream  habitat  or  stable  stretches  of 
shallow  riffles  in  somewhat  larger  rivers  like  the 
Duck.  Stansbery  (1976)  comments  that  "Pegias 
fabula  appears  to  be  a  rare  Cumberlandian 
species  characteristic  of  stream  conditions  near, 
but  not  quite  in,  the  uppermost  headwaters." 

The  three-ridge,  Amblema  plicata,  and  purple 
warty-back,  Cyclonaias  tuberculata,  are  two  of 
the  more  common  species  inhabiting  the  Duck 
River  today.  The  27  valves  of  A.  plicata  com- 
prised only  3.5%  of  the  total  sample  but  were  of 
interest  in  that  they  exhibited  characteristics 
(small  and  compressed)  of  those  inhabiting  a 
headwaters  or  small  stream  habitat.  Only  two 
valves  of  the  latter  species  were  recovered  in 
the  cave.  Lexingtonia  dolabelloides,  a  species 
fairly  well  represented  in  the  Cheek  Bend  Cave 
naiad  assemblage  (ca.  6%  of  the  identified 
valves),  still  occurs  in  the  Duck  River,  one  of  the 
few  remaining  rivers  still  supporting  viable 
populations  of  this  mussel.  Quad^-ula  inter- 
media is  another  species  of  interest  in  that  it 
was  present  in  the  middle  stretches  of  the  Duck 
River  in  prehistoric  times  (2  valves  from  Cheek 
Bend  Cave)  and  continued  to  survive  until  pre- 
sent (Ahlstedt,  1981)  but  at  very  low  population 
levels.  It  was  apparently  never  a  common  shell 
in  the  Duck  River  as  evidenced  by  the  paucity  of 
specimens  recovered  at  Cheek  Bend  Cave  and  at 
the  Hayes  site  where  only  five  shells  of  Q.  in- 
termedia were  identified  out  of  a  sample  of 
3,870  valves  (Turner.nd). 

Anomalous  Valves 

During  the  identification  process  it  was 
necessary   to   compare   certain   archaeological 


138    THE  NAUTILUS 


October  31,  1986 


Vol.  100  (4) 


Table  1.   Freshwater  mussels  Identified  from  Cheek  Bend  Cave.  Maury  County.  Tenness. 
Percent  of  each  species  within  Strata  and  for  Total  Valves  listed  In  parentheses  { 


Amblema  pllcata  (Say.  1817) 
Three-Rld^e 

Fusconala  barnestana  (Lea.  1838) 
Tennessee  Plgtoe 

Lexington la  dolabelloldes  (Lea,  1840) 
Slabslded  Pearlyoussel 

F.  bameslana  and/or  L.  dolabelloldes 


Quadrula  CvllQ(^l.ca  (Say,  1817) 
Rabbitsfoot 


VII 

13 
(4.19) 

5 

(1.61) 

10 
(3.23) 

2 
(.65) 


7 
(12.28) 

2 
(.79) 

4 
(3.13) 

2 
(3.5) 

4 
(1.57) 

2 
(1.56) 

A 
(7.02) 

23 
(9.06) 

1 
(.39) 

10 
(7.81) 

Total 
Valves 

27 
(3.55) 

13 
(1.71) 

47 
(6.18) 

3 
(.39) 


Quadrula  InCenpedla  (Conrad,  1836) 
Cumberland  Monkeyface 

Cyclonalas  tuberculata  (Raflnesque,  1820) 
Purple  Wartyback 


Elliptlo  dllatata  (Raflnesque.  1820) 
Spike 

Hemlstena  lata  (Raflnesque.  1820) 
Cracking  Pearlymussel 

Pleurobema  ovlforme  (Conrad.  1834) 


Tennessee  Clubshell 


Alasmldonta  virldla  (Raflnesque,  1820) 
Slippershell 


Peglas  Fabula  (Lea.  1836) 

Llttle-wlnged  Pearly  Mussel 


Lasaigona  costata  (Rafineaque,  1820) 
Fluted  Shell 


SCrophitus  undulatu! 
Squawfoot 


Actlnonalas  pectorosa  (Conrad,  1834) 
Pheasant she 11 

Toxolasma  cyllndrellus  (Lea.  1868) 
Pale  Lllllput 

Toxolasma  llvldus  (Raflnesque,  1831) 
Purple  Lllllput 

Eploblasma  capsaeformls  (Lea.  1834) 

and /or  E.  fjgrentlna  (Lea.  1857) 
Oyster  HusseT  i/or  Tan  Rlffleshell 

Eploblasma  brevldens  (Lea,  1834) 
Cumberland  Combshell 


Lanpsllls  fasclola  (Raflnesque,  18 
Wavy-rayed  Lampmussel 

Lampsllls  ovata  (Say.  1817) 
Pocket book 

Lemiox  rimosus  (Raflnesque.  1831) 
Blrdwlng  Pearlymussel 

Medlonldus  conradlcus  (Lea,  1834) 
Cumberland  Moccasin 

Potamllus  alaCus  (Say,  1817) 
Pink  Heelsplitter 


1 
(8.33) 


(.65) 

10 
(3.23) 

4 
(1.29) 

1 
(.32) 

2 

(.65) 


(.97) 

1 

(8.33) 

17 
(5.48) 

1 
(8.33) 

5 

(1.61) 

4 
(33.33) 

49 
(15.81) 

1 
(.32) 

(3.51) 

1 

(1.75) 

10 
(17.54) 


1 

(.39) 

1 
(.78) 

2 
(.26) 

1 
(.78) 

2 
(.26) 

43 
(16.93) 

9 

(7.03) 

90 
(11.83) 

1 
(.39) 

1 
(.78) 

2 
(.26) 

1 
(.39) 

4 
(.53) 

1 

(.39) 

4 
(3.13) 

8 
(1.05) 

1 
(.78) 

11 
(1.45) 

2 
(1.56) 

6 
(.79) 

1 
(.13) 

4 
(1.57) 

8 
(6.25) 

15 
(1.97) 

1 
(.39) 

5 
(.66) 

1 
(.39) 

(.13) 

40 
(15.75) 

16 

(12.50) 

150 
(19.71) 

1 
(.39) 

1 
(.13) 

10 
(3.94) 

1 

(.78) 

14 
(1.84) 

(4.69) 

23 

(17.97) 


13 
(1.71) 


100 

(13.14) 


Vlllosa  Iris  (L«a,  1830) 
Rainbow 


Vlllosa  taenlata  (Conrad.  1834) 
Painted  Creekshell 


Vlllosa  vanuxemensls  (Lea,  1838) 
Mountain  Creekshell 


Vlllosa  spp.  (Eploblasma?) 


Ptychobranchus  faaciolare  (Raflnesqui 
Kldneyshell 


Ptychobranchus  sub ten cum  (Say, 
Fluted  Kldneyshell 


17 
(5.48) 


(1.29) 

10 
(3.12) 


2 
(3.51) 

3 
(1.18) 

2 
(1.56) 

11 
(1.45) 

1 
(1.75) 

35 

(13.76) 

20 
(15.63) 

73 
(9.59) 

2 
(3.51) 

1 

(.39) 

1 
(.78) 

8 
(1.05) 

1 
(1.75) 

8 
(3,15) 

4 
(3.13) 

24 
(3.15) 

1 
(.39) 

1 
(.13) 

4 
(7.02) 

58 
(22.83) 

12 
(9.38) 

107 
(14.06) 

Totals 

(Identified  valves) 


Totals 
(Unidentified  valves) 


(99.97) 
17 


310 
(99.90) 


57       254 
(99.99)    (99.96) 


128        761 
(100.01)    (100.00) 


Totals 
(All  Valves) 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    139 


specimens,  particularly  incomplete  valves  and 
those  of  Villoma  spp.  and  Epioblnsma  spp.,  with 
fresh  material  using  a  binocular  scope.  Under 
magnification,  a  variety  of  anomalous  struc- 
tures or  conditions  were  noted  that,  in  addition 
to  making  specific  determinations  difficult  or 
impossible,  pose  some  interesting  questions. 
These  anomalies  consisted  primarily  of  deep  pits 
along,  under  and/or  between  the  lateral  and 
pseudocardinal  teeth  and  the  development  of 
supernumerary  "teeth"  or  projections  between 
or  adjacent  to  the  pseudocardinal  teeth  (Fig.  3). 
Less  than  3%  of  the  valves  exhibited  one  or 
more  of  these  anomalies;  the  majority  occurred 
in  small,  thin-shelled  individuals.  Possibly  these 
conditions  were  the  result  of  some  trauma  or 
perhaps  parasites,  but  whatever  the  cause(s). 


F^IG.  3.   Examples  of  anomalous  naiad  valves  from  Cheek 
Bend  Cave  that  exhibit  pitting  and  nacre  deformities. 


such  abnormalities  have  not  been  observed  in 
other  archaeological  naiad  material  from  pre- 
historic sites  along  the  Duck  River. 

General  Discussion 

Ortmann  (1924:19),  in  attempting  to  evaluate 
the  zoogeographical  affinities  of  the  Duck  River 
naiad  assemblage,  made  the  following  signifi- 
cant observations:  "Duck  River  being  located 
between  Cumberland  River  in  Kentucky  and 
Tennessee,  and  Tennessee  River  in  northern 
Alabama,  might  be  expected  to  contain  a  similar 
fauna  to  these  two  systems,  a  fauna  which  is 
known  to  contain  peculiar  elements,  not  found 
outside  of  this  region  in  the  Central  basin 
(Mississippi  and  Ohio  drainages),  which  might  be 
called  'Cumberlandian'  types  (from  the 
'Cumberland  subregion')."  On  the  basis  of  his 
studies  of  the  Duck  River  mussel  species  and 
those  of  other  rivers  of  the  southeastern  region, 
he  concluded  (Ortmann  1924:61)  that  "It  ap- 
pears that  the  Cumberlandian  fauna  is  the 
original  fauna  of  Duck  River,  while  the  interior- 
basin-fauna  is  a  later  invasion  of  the  river,  com- 
ing up  from  the  lower  parts;  yet  important 
elements  of  the  interior  fauna  have  not  yet 
reached  Duck  River.  In  the  lower  Duck,  at  Cen- 
treville,  and  probably  farther  down,  the  interior 
fauna  alone  is  present,  and  this  part  does  not 
belong  to  the  Cumberland  region.  Duck  River 
originally  was  more  directly  connected  with  the 
Cumberland  and  Tennessee,  and,  at  that  time,  it 
was  a  rather  small  river."  Approximately  65%  of 
the  species  represented  in  the  Cheek  Bend  naiad 
assemblage  are  Cumberlandian  as  defined  by 
Ortmann,  and  they  reflect  a  headwaters  or  small 
stream  habitat.  Several  other  species  (e.g.  E. 
dUatata,  A.  plicnta)  present  in  the  cave  with 
Mississippian  or  Interior  Basin  affinities  exhibit 
small  stream  shell  characters,  that  is,  com- 
pressed, thin-shelled  small  valves.  Therefore, 
the  naiad  assemblage  recovered  from  the  Cheek 
Bend  Cave  excavations  and  the  occurrence  of 
certain  species  such  as  P.  fahula  from  this  cave 
and  other  aboriginal  sites  in  Maury,  Marshall 
and  Bedford  counties  suggest  that  the  Duck 
River  in  Middle  Tennessee  was  swift  and 
shallow  with  riffles  and  a  probable  substrate  of 
sand  and  small  gravel  for  extended  periods  be- 
tween 7,000  and  1,000  BP. 


140    THE  NAUTILUS 


October  31.  1986 


Vol.  100(4) 


Acknowledgments 

We  would  like  to  express  our  appreciation  to 
Terry  Faulkner  for  preparing  Fig.  1  and  to 
Betty  W.  Creech  for  typing  the  manuscript. 
Special  thanks  are  extended  to  W.  Miles  Wright 
for  photographing  the  specimens  in  Fig.  3  and 
the  preparation  of  Figs.  2  and  3.  We  gratefully 
acknowledge  the  Tennessee  Valley  Authority 
under  whose  auspices  (contracts  No.  TVA  TV- 
49244A  and  TVA  TV-53013A)  the  excavation  of 
Cheek  Bend  Cave,  plus  the  analysis  of  the  faunal 
materials  recovered,  were  made  possible. 

LITERATURE  CITED 

Ahlstedt,  S.  A.  1981.  The  molluscan  fauna  of  the  Duck  River 
between  Normandy  and  Columbia  dams  in  central  Tennes- 
see. Bull.  Amer.  Malacol.  Union  for  1980.  pp.  60-62. 

Clarke,  A.  H.,  Jr.  1981.  The  Tribe  Alasmidontini  (Unionidae: 
Anodontinae),  Part  I:  Pegias.  Alastnidonta.  and  Ar«rf«i.s. 
Smithsonian  Contributions  to  Zoology  No.  326.  101  pp. 

Hinkley,  A.  A.  and  P.  Marsh.  1885.  Lis?  of  shells  collected  in 
central  Tennessee.  Published  privately.  Aledo,  Illinois. 
10  pp. 

Hofman.  J.  L.  nd.  Ervin;  A  mid  Holocene  shell  midden  on  the 
Duck  River  in  the  Nashville  Basin  of  Tennessee.  Manu- 
script on  file,  Department  of  Anthropology,  University  of 
Tennessee,  Knoxville,  TN. 

Isom,  Billy  G.  and  Paul  Yokley,  Jr.  1968.  The  mussel  fauna 
of  Duck  River  in  Tennessee,  1965.  Airie?'ican  Midland 
Naturalist  80(l);34-42. 

Klippel,  W.  E.  and  P.  W.  Parmalee.  1982.  Diachronic  varia- 


tion in  insectivores  from  Cheek  Bend  Cave  and  environ- 
mental change  in  the  Midsouth.  Paleobiology  8(4):447-458. 

O'Hare,  C.  R.  nd.  Goatcliff  and  McCoUum  rock.shelters: 
analysis  of  cultural  and  non-cultural  processes  operating 
on  the  formation  of  two  faunal  samples.  MA  thesis  in 
preparation.  Department  of  Anthropology.  University  of 
Tennessee,  Knoxville,  TN. 

Ortmann,  A.  E.  1924.  The  naiad-fauna  of  Duck  River  in  Ten- 
nessee. American  Midland  Naturalist  9(2):  18-62. 

Parmalee,  P.  W.  and  W.  E.  Klippel.  1974.  Freshwater 
mussels  as  a  prehistoric  food  resource.  American  Anti- 
quity 39(3):421-434. 

1981.  A  late  Pleistocene  population  of  the  pocket 

gopher,  Geomys  cf.  bursarus,  in  the  Nashville  Basin,  Ten- 
nessee. Jour.  Mammalogy  62(4):831-835. 

Robison,  N.  D.  nd.  An  analysis  and  interpretation  of  the 
faunal  remains  from  eight  late  Middle  Woodland  Owl 
Hollow  phase  sites  in  Coffee,  Franklin  and  Bedford  coun- 
ties, Tennessee.  Ph.D  dissertation  in  preparation,  Depart- 
ment of  Anthropology,  University  of  Tennessee,  Knox- 
ville, TN. 

Stansbery,  D.  H.  1976.  Status  of  endangered  fluviatile 
mollusks  in  central  North  America:  Pegias  fabula  (Lea, 
1838).  Report  prepared  for  the  U.S.  Department  of  the  In- 
terior, Fish  &  Wildlife  Service  and  the  Bureau  of  Sport 
Fisheries  &  Wildlife,  Washington,  D.C.  8  pp. 

Turner,  W.  B.  1984.  A  preliminary  analysis  of  prehistoric 
fresh-water  bivalve  species  composition  changes  from  ca. 
7500  to  2500  B.P.:  a  view  from  the  central  Duck  River 
drainage.  Manuscript  on  file.  Department  of  Anthro- 
pology. University  of  Tennessee,  Knoxville,  TN. 

van  der  Schalie,  H.  1973.  The  mollusks  of  the  Duck  River 
drainage  in  central  Tennessee.  Sterkiana  52:45-56. 


THE  FATE  OF  PELECYPOD  FAMILIES,  SUBFAMILIES,  AND  TRIBES 
DURING  AND  AFTER  THE  CRETACEOUS  PERIOD 

David  Nicol 

Box  14376,  University  Station, 
Gainesville,  FL  32604 


ABSTRACT 
Of  the  lU  surveyed  supergeneric  groups  of  pelecypods  that  occur  in  the 
Cretaceous,  38,  or  33%,  are  extinct.  In  the  Early  Cretaceous  one  group  became  ex- 
tinct every  9,300,000  years.  During  the  Late,  but  not  latest.  Cretaceous,  one  group 
became  extinct  every  2,333,000  years.  In  the  Maastrichtian  Stage  (latest 
Cretaceous),  the  extinction  rate  increased  to  one  group  every  470.000  years.  Only 
seven  groups  became  extinct  during  the  Cenozoic:  four  during  the  Eocene  and 
three  during  the  Miocene.  There  was  only  one  extinction  of  a  supergeneric  group 
every  9,500,000  years  during  the  entire  Cenozoic,  which  is  comparable  to  the  ex- 
tinction rate  during  the  Early  Cretaceous.  The  rate  of  extinction  of  supergeneric 
groups  of  pelecypods  increased  rapidly  in  the  Late  Cretaceous  and  reached  a 
climax  at  the  end  of  that  geologic  period. 


The  number  of  families,  subfamilies,  tribes 
and  groups  of  pelecypods  that  undoubtedly  had 


one  or  more  genera  living  during  the  Cretaceous 
Period,  as  reviewed  in  the  Treatise  on  Inverte- 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    141 


brate  Paleontology  (1969,  1971),  is  124.  In  each 
of  these  supergeneric  categories  the  genera  and 
subgenera  were  tallied  in  each  geologic  Period 
and  Epoch.  In  a  few  instances,  more  current 
data  were  used  when  it  was  available  (Keen's, 
1980,  latest  review  of  the  Cardiidae),  and  a  few 
of  the  geologic  ranges  of  other  supergeneric 
groups  have  been  brought  up  to  date.  Ten  of  the 
124  supergeneric  groups  were  excluded  because 
of  insufficient  geologic  data,  as,  for  example, 
the  Solemyidae,  Manzanellidae,  Dimyidae,  and 
Sportellidae. 

The  supergeneric  groups  that  lived  during  the 
Cretaceous  are,  commonly,  not  ancient  line- 
ages. Only  the  Nuculanidae  and  Malletiidae  can 
be  traced  back  to  the  Ordovician,  and  they  are 
as  diverse  today  as  they  have  ever  been.  Fifty 
three  supergeneric  groups  (47%)  appear  in  the 
Cretaceous.  Some  of  these  are  short-lived 
groups  but  many  others  first  appeared  in  the 
Cretaceous  and  are  presently  represented  by 
many  genera  and  species.  By  the  end  of  the 
Cretaceous,  most  of  the  basic  adaptive  radiation 
of  the  pelecypods  had  occurred.  Of  the  114 
supergeneric  categories  considered  herein,  90% 
first  appeared  in  the  Mesozoic.  Stanley's  graph 
(1968,  p.  215)  depicts  the  great  Mesozoic 
radiation. 

The  114  supergeneric  groups  can  be  placed  in 
seven  categories  based  on  the  distribution  of 
their  genera  and  subgenera  from  the 
Cretaceous  to  the  Recent. 

1.  Many  supergeneric  groups  were  relatively 
unaffected  by  the  mass  extinctions  at  the  end  of 
the  Cretaceous  and  remained  as  diverse  in  the 
Cenozoic  or  increased  in  diversity  during  that 
Era.  In  this  first  category  there  are  46  super- 
generic  groups  or  40%  of  the  total  of  114. 
Although  some  genera  became  extinct  during 
the  Cretaceous,  they  were  quickly  replaced  by 
new  genera  arising  in  the  Cenozoic.  Almost  62% 
of  these  supergeneric  groups  began  in  the 
Cretaceous  and  87%  began  after  the  Paleozoic. 
Those  groups  first  appearing  in  the  Cretaceous 
are  commonly  represented  by  one  or  two 
genera.  In  other  words,  they  are  comparatively 
young  supergeneric  groups.  The  protobranch 
families  Nuculidae,  Malletiidae,  and  Nuculani- 
dae are  exceptional  in  that  they  range  well  back 
into  the  Paleozoic  and  were  not  affected  greatly 
by  extinction  at  the  end  of  the  Mesozoic. 


2.  Some  supergeneric  groups  were  affected 
by  Cretaceous  extinction  but  later  recovered 
and  became  most  diverse  in  the  Neogene.  There 
are  15  supergeneric  groups  that  have  this 
distribution  in  diversity,  which  comprise  about 
13%  of  the  total.  In  this  category  only  six  or 
40%  began  in  the  Cretaceous  but  93%  began  in 
the  Mesozoic.  In  general,  the  families  are  longer 
ranging  than  those  that  were  little  affected  by 
the  Cretaceous  extinctions.  Some  notable  ex- 
amples of  supergeneric  groups  in  this  category 
of  diversity  are  the  Pitarinae,  Tellininae,  Car- 
diinae,  Protocardiinae,  and  Mactrinae. 

3.  Some  supergeneric  groups  were  affected 
by  Cretaceous  extinction  but  continued  on  to  the 
Recent  with  a  reduction  in  diversity.  In  this 
category  are  included  15  supergeneric  groups  or 
approximately  13%  of  the  total.  These  super- 
generic  groups  are  still  longer  ranging  than 
those  in  the  first  two  categories  and  only  four, 
or  27%,  began  in  the  Cretaceous.  However,  13, 
or  87%,  had  their  inception  in  the  Mesozoic. 
There  are  some  notable  groups  that  were 
adversely  affected  by  the  Cretaceous  wave  of 
extinction,  including  the  Trigoniidae,  which  has 
survived  to  the  present  only  in  the  Australian 
region.  The  Pholadomyidae  is  another  group 
that  is  nearly  extinct  after  its  great  diversity  in 
the  Mesozoic.  Other  relict  groups  at  present  are 
the  Grammatodontidae,  Cucullaeidae,  Fim- 
briidae,  and  Arcticidae.  All  of  these  families 
were  much  more  diverse  and  widespread  during 
the  Cretaceous. 

Of  the  114  supergeneric  groups  studied,  38,  or 
33%  are  extinct.  They  can  be  grouped  into  four 
categories. 

4.  A  few  supergeneric  groups  became  extinct 
during  the  Early  Cretaceous  (before  the 
Cenomanian).  There  were  only  five  families  or 
about  4.4%  of  the  supergeneric  groups  that 
became  extinct  during  this  span  of  time.  Eighty 
percent  of  this  small  group  are  confined  to  the 
Mesozoic,  which  includes  the  Neomiodontidae,  a 
small  family  that  lived  in  brackish  and  fresh 
water. 

5.  There  are  nine  supergeneric  groups  of 
pelecypods  that  became  extinct  in  the  Late 
Cretaceous  but  before  the  Maastrichtian.  This 
category  constitutes  7.9%  of  the  total  number  of 
supergeneric  groups.  Four  of  these  are  small 
and  short-lived  families  and  subfamilies  that 


14:^    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


were  confined  to  the  Cretaceous  Period.  One  of 
these  short-lived  families,  the  Trigonioididae, 
lived  in  fresh  and  brackish  water.  I  have  placed 
the  Mactromyidae  in  this  category  because 
Bathycorbis  Iredale  is  most  likely  an  aberrant 
venerid  and  Dockery  (1982,  p.  72)  has  proved 
that  Cordiula  Meyer  was  based  on  a  young 
specimen  of  Nemocardiurn.  The  Mactromyidae 
is  found  primarily  in  the  Jurassic  and 
Cretaceous. 

6.  Seventeen  supergeneric  groups,  about  15% 
of  the  total,  became  extinct  at  the  end  of  the 
Cretaceous  during  the  Maastrichtian  Stage. 
Some  of  the  supergeneric  groups  that  became 
extinct  at  this  time  were  large-sized,  diverse, 
and  common  in  strata  of  Cretaceous  age.  These 
include  the  rudist  families  Hippuritidae, 
Radiolitidae,  Caprinidae,  Monopleuridae,  and 
Requiniidae.  It  also  includes  the  Inoceramidae 
and  Exogyrini.  Seven  of  these  seventeen  groups 
(41%)  were  confined  to  the  Cretaceous  Period, 
and  14  (82%)  were  found  only  in  the  Mesozoic. 
The  great  extinction  at  the  end  of  the 
Cretaceous  did  not  greatly  affect  most  of  the 
older  supergeneric  groups. 

7.  Seven  supergeneric  groups  (6.1%)  survived 
the  Cretaceous  but  became  extinct  in  the  Ter- 
tiary-four in  the  Eocene  and  three  in  the 
Miocene.  Except  for  the  Bakevelliidae  and 
Eriphylinae,  they  consist  of  small  groups  (no 
more  than  four  genera),  and  all  hut  the  Bakevel- 
liidae and  Eriphylinae  originated  in  the 
Cretaceous  Period.  It  is  known  that  extinction 
rates  increased  near  the  end  of  the  Eocene 
Epoch,  as  in  the  planktonic  Foraminiferida,  and 
for  this  reason  it  is  not  surprising  that  a  few 
supergeneric  groups  of  pelecypods  disappeared 
at  this  time.  Two  of  the  three  extinctions  of 
supergeneric  groups  that  occurred  in  the 
Miocene  were  the  Gryphaeostreinae  and  the 
Flemingostreinae  (Ostracea)  and  may  have  been 
caused  by  oncoming  glaciation  at  the  end  of  the 
Miocene  (Nicol,  1984). 

Fresh-water  pelecypods  were  somewhat  af- 
fected by  the  Cretaceous  extinctions.  Besides 
the  two  previously  mentioned  short-lived  groups 
that  became  extinct  in  the  Cretaceous,  the  Cor- 
biculidae  and  Unioninae  lost  some  genera  and 
subgenera  at  the  end  of  the  Mesozoic. 

In  Table  1  I  have  summarized  the  rate  of  ex- 
tinction  of  the  supergeneric  groups  of  pele- 


TABLE  1.  Extinction  rates  of  families,  subfamilies,  and 
tribes  of  pelecypods  during  the  Cretaceous  and  Cenozoic. 

Stratigraphic  time  units  Number  of  Extinction 

and  time    in  years  extinctions         rate   in  years 

1.  Early  Cretaceous 
(Berriasian-Albian) 

i.0, 500,000  5  9,300,000 

2.  Late  but  not  latest 
Cretaceous  (Cenomanian- 

Campanian)  21,000,000  9  2,333,000 

3.  Latest  Cretaceous 

(Maastrichtian)  8,000,000      17  470,000 

4.  Late  Cretaceous 
(Cenomanian-Maastrichtian) 

29,000,000  20  1,115,000 

5.  Total  Cretaceous 

75,500,000  31  2,1.35,000 

0,  Paleocene-Eocene 

30,000,000  4  7,500,000 

7.  Oligocene-Recent 

30,500,000  3  12,170,000 

3.  Total  Cenozoic 

00,500,000  7  9,500,000 

9.  Total  of  Cretaceous 

and  Cenozoic  142,000,000       38  3,740,000 

cypods  in  years  throughout  the  Cretaceous  and 
Cenozoic.  The  actual  time  spans  of  the  various 
stratigraphic  units  were  taken  from  Palmer, 
1983.  The  extinction  rate  was  low  in  the  Early 
Cretaceous,  but  beginning  with  the  Cenoma- 
nian,  which  is  the  earliest  Stage  of  the  Late 
Cretaceous,  the  extinction  rate  increased  and 
continued  into  the  Campanian  Stage.  This  latter 
rate  is  a  fourfold  increase  over  the  Early 
Cretaceous  rate  of  extinction.  The  Maastrich- 
tian Stage  at  the  end  of  the  Cretaceous  has  the 
highest  rate  of  extinction,  being  about  five  times 
faster  than  the  previous  figure  in  the  rest  of  the 
Late  Cretaceous,  and  it  is  about  20  times  faster 
than  the  rate  in  the  Early  Cretaceous.  Further- 
more, the  extinction  rate  in  all  of  the  Late 
Cretaceous  was  more  than  eight  times  more 
rapid  than  it  was  in  the  Early  Cretaceous.  These 
extinction  rates  do  not  appear  to  be  an  artifact 
of  the  geologic  record.  Cretaceous  strata,  par- 
ticularly that  of  a  shallow-water  marine  environ- 
ment,  are   exceedingly   widespread   and   have 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    143 


been  thoroughly  studied  by  invertebrate  paleon- 
tologists. Evolution  increased  greatly  in  rate 
during  the  Late  Cretaceous,  a  time  span  of 
29,000,000  years.  This  rate  of  evolution  in- 
creased and  reached  a  clima.\  in  the  latest 
Cretaceous  Maastrichtian  Stage,  which  spans 
only  8,000,000  years.  For  example,  three  of  the 
supergeneric  groups  that  arose  in  the  Late 
Cretaceous  also  became  extinct  during  that  part 
of  the  Cretaceous  Period.  Apparently  new 
species,  genera,  and  families  arose  with  increas- 
ing rapidity  during  the  Late  Cretaceous  as 
evinced  by  the  rudists  (Jones  and  Nicol,  in  press, 
and  Nicol,  in  press).  The  Inoceramidae  also 
evolved  rapidly  during  the  Late  Cretaceous. 
One  may  ask,  did  the  widespread  warm  shallow 
seas  that  covered  the  continents  during  the  Late 
Cretaceous  cause  evolutionary  rates  to  increase 
greatly? 


LITERATURE  CITED 

Dockery,  D.  T.,  III.  1982.  Lower  Oligocene  Bivalvia  of  the 
Vicksburg  Group  in  Mississippi.  A/).s-.s-(.s.s-(p/))  Bureau  of 
Geology.  Bull.  123,  261  p. 

Jones,  D.  S.  and  D.  Nicol.  Origination,  survivorship,  and  ex- 
tinction of  rudist  taxa.  Jour.  Paleontology  60:107-115. 

Keen.  A.  M.  1980.  The  pelecypod  family  Cardiidae:  A  tax- 
ononiic  summary.  Tulnne  Studief:  in  Geology  and  Paleon- 
tology 16:1-40. 

Moore,  R.  C.  (ed.).  1969,  1971.  Treatise  on  Invertebrate 
Paleontology.  Part  N,  Mollusca  6,  Bivalvia,  Vols.  1-2 
(1969).  Vol.  3  (1971).  Univ.  Kansas  Press,  Lawrence,  p. 
N1-N1224. 

Nicol,  D.  1984.  Critique  on  Stenzel's  book  on  the  Ostracea. 
The  Nautilus  98:123-126. 

Some  aspects  of  the  evolution  of  the  rudist  pele- 

cypods.  The  Nautilus  100(2):69-71. 

Palmer,  A.  R.  1983.  The  decade  of  North  American  geology 
1983  geologic  time  scale.  Geology  ll:.503-.504. 

Stanley,  S.  M.  1968.  Post-Paleozoic  adaptive  radiation  of  in- 
faunal  bivalve  molluscs -A  consequence  of  mantle  fusion 
and  siphon  formation.  Jour.  Paleontology  42:214-229. 


FRESHWATER  AND  TERRESTRIAL  SNAILS  OF 
SAINT  LUCIA,  WEST  INDIES^ 

Emile  A.  Malek 

Department  of  Tropical  Medicine 

Tulane  University  Medical  Center 

1430  Tulane  Avenue,  New  Orleans,  LA  70112 

ABSTRACT 

Surveys  were  made  of  the  Jreshwater  and  terrestrial  snails  and  slugs  of  Saint 
Lucia,  one  of  the  Windward  Islands  of  the  Lesser  Antilles.  Ten  species  of  7 
families  of  freshwater  snails,  9  species  of  7  families  of  terrestrial  snails,  and  2 
species  of  one  family  of  slugs  were  encountered.  The  freshwater  snail  fauna 
inhabit  all  types  of  freshwater  habitats  available,  i.e.,  rivers,  br-ooks  (ravines), 
stream  embankments,  flood  plains  and  swamps  adjacent  to  rivers,  drainage 
ditches  and  ponds.  Only  one  species  of  Biomphalaria,  B.  glabrata,  was  found, 
although  other  species  of  this  genus  occur  on  some  other  neighboring  islands,  alone 
or  together  with  B.  glabrata.  The  majority  of  the  land  srmil  species  were  repre- 
sented in  the  capital  city,  Castries,  and  the  nearby  hills  and  coastal  lands.  Cer- 
tain species  of  land  snails  were  associated  with  limestone  deposits  and  others, 
although  inhabiting  the  same  areas,  were  also  found  in  the  igneous  mountain 
highlands  and  in  Jbrests.  A  scarcity  of  land  operculates  was  observed;  only  one 
species  was  found. 


There  is  some  old  information  on  the  mollus- 


'Contribution  from  the  World  Health  Organization  Colla- 
borating Center  for  Applied  Medical  Malacology,  Tulane 
University. 


can  fauna  of  Saint  Lucia.  Bland  (1861),  E.  A. 
Smith  (1889,  1895)  and  E.A.  Smith  and  Feilden 
(1891)  listed  land  snails  and  a  few  freshwater 
snails  of  Saint  Lucia  and  some  neighboring 
islands.  The  author  (1965),  in  an  abstract  form. 


144    THE  NAUTILUS 


October  31,  1986 


Vol.  100  (4) 


enumerated  and  discussed  briefly  the  fresh- 
water and  land  snails  based  on  three  trips  to 
Saint  Lucia  for  which  reports  to  the  Pan 
American  Health  Organization  were  prepared 
(Malek,  1962,  1963).  The  snails  were  later 
(Malek,  1980)  illustrated  to  emphasize  shell 
characteristics.  Jordan  (1985)  in  a  brief  appen- 
dix included  only  a  short  list  of  the  freshwater 
and  terrestrial  snails  on  Saint  Lucia,  some  of 
which  were  identified  by  the  author. 

The  present  report  was  prepared  in  view  of 
the  significance  of  the  molluscan  fauna  in 
research  on  snail-transmitted  diseases  on  the 
island,  especially  schistosomiasis  and  fas- 
cioliasis,  and  the  reports  in  recent  years  on 
freshwater  snails  in  other  Antillean  islands  such 
as  Guadeloupe  (Pointier,  1974),  Haiti  (Robart  et 
al.,  1976),  Martinique  (Guyard  and  Pointier, 
1979),  Guadeloupe,  Dominica  and  Martinique 
(Starmuhlner,  1985)  and  the  Dominican 
Republic  (Gomez  et  al.,  1986). 

The  Island  of  Saint  Lucia 

Saint  Lucia,  one  of  the  Windward  Islands 
located  between  Martinique  and  Saint  Vincent, 
is  a  small  island  27  by  14  miles  and  is  about  233 
square  miles  in  area.  The  middle  part  is  moun- 
tainous; there  is  a  longitudinal  main  ridge, 
almost  median,  with  other  mountains  running 
down  to  the  coast  on  either  side.  Between  the 
offshoots  of  the  main  mountain  ridge  are  broad, 
flat  valleys  which  spread  out  toward  the  sea. 
Small  rivers,  measuring  only  7  to  12  feet  in 
width  in  most  parts,  and  only  a  few  feet  in 
depth,  run  from  the  mountain  heights  through 
the  valleys.  Most  rivers  form  separate  drainage 
systems  that  are  subject  to  frequent  flooding. 

Rainfall  over  most  of  the  island  averages  80 
inches  per  year,  but  with  only  40  inches  on  the 
southern  and  northern  tips.  The  highest  rainfall 
occurs  from  May  through  October,  with  peaks  in 
August  and  September.  Remnants  of  past 
volcanic  activity  are  still  noticeable  in  the 
sulphur  area  near  Soufriere.  The  river  beds  have 
big  rocks,  seemingly  of  volcanic  origin.  Also  in 
the  geologic  history  certain  areas  were  elevated, 
and  during  the  Cretaceous  period,  deposition  of 
limestone  took  place  on  several  parts  of  the 
island,  which  support  large  numbers  of  the 
molluscan  species. 


Results 

Ten  species  belonging  to  7  families  of 
freshwater  snails,  9  species  belonging  to  7 
families  of  terrestrial  snails,  and  2  species  of  one 
family  of  slugs  were  encountered  in  this  survey 
(Table  1).  Of  the  freshwater  snails  the  Planorbi- 
dae  was  represented  by  3  species,  the  Neritidae 
by  2  species  and  the  Lymnaeidae,  Physidae,  An- 

TABLE  1 .  Families  and  species  of  gastropods  and  their  loca- 
tions in  Saint  Lucia. 

FRESHWATER 

Planorbidae 

Biomphala ria  glnbrata 
Rivers:  Castries,  in  the  Entrepot  area;  Derniere,  north  of 
Dennery;  Esperance,  near  Fond  St.  Jacques;  Bance,  near 
bridge  south  of  school;  Choc;  Raveneau,  at  Delcer.  Brooks 
(ravines):  Ti  Rocher;  Trou  Macis  near  Bance;  above  fall  at 
Sulphur  Springs.  River  and  brook  embankments:  Ravine 
Poisson  at  Ravine  Poisson;  Augier  River.  Flood  plains  or 
swamps  off  rivers:  Vieu.x  Fort  north  of  Joyeox;  Canaries; 
Augier.  Drainage  ditches:  Banana  fields  near  Marquis  and 
in  Cul  de  Sac  Valley.  Ponds:  At  fall  near  Sulphur  Springs; 
at  Fond  St.  Jacques;  at  Derniere  Fort.  Concrete-lined 
channels:  At  Ferrand  Estate. 

Drepanotrema  depressissimum. 
Pond  at  entrance  of  Union  Agricultural  Station;  pond  in 
Bonne  Terre  area;  pond  in  Ville  Boutille;  pond  near 
Monchy  school;  drainage  ditch  in  Marquis  Estate;  Ti 
Rocher  brook;  swamp  by  road  between  Vieux  Fort  and 
Labourie. 

D repanotrema  I uctiiu m 
Pond  near  Monchy  school;  pond  in  Bonne  Terre  area;  pond 
in  Ville  Boutille. 

Lymnaeidae 

Fossaria  cubensis 
Ditches  in  banana  fields,  Cul  de  Sac  Valley;  along  bank  of 
river  near  LInion  Agricultural  Station;  Ferrand  Estate. 

Physidae 

Phusn  ruheiisis 
Derniere  River;  embankment  along  Poisson  brook;  Mar- 
quis River;  Bagatelle  River  in  the  Marchand  area;  pond  in 
Bonne  Terre  area;  pond  at  Ville  Boutille;  pond  in  Monchy 
area;  brook  joining  Vieux  Fort  River  north  of  Joyeax. 

Ancylidae 

Gundlachia  radiata 
Pond   near  Sulphur  Springs;   Derniere   River;   Castries 
River  in  Marchand  area;  Choc  River;  ditches  in  banana 
fields,  Marquis  Estate. 

Hydrobiidae 

Pyrgopkonis  parmtlus 
Brook,  near  Fond  St.  .Jacques;  brook  near  Joyeux;  brook  in 
Cul  de  Sac  Valley. 

Anipullaridae 

Atiipulldnii  ghiiica 

Derniere  River;  Marquis  River;  Bonne  Terre  pond;  pond  at 
entrance  of  Union  Agricultural  Station. 

Neritidae 

Neritina  punctulata  and  N.  virginea 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    145 


Castries  River;  Marquis  River;  Choc  River 
TERRESTRIAL 
Helicinidae 
Helicimi  fasriata 

Ferrand  Estate,  on  banana  plants  and  weeds  along  drain- 
age ditches. 
Suceineidae 
Omalonyx  guadeloupensis 

Banks  of  pond  at  entrance  of  Union  Agricultural  Station; 

Cul  de  Sac  Valley;  along  ditches  in  banana  fields. 
Succinea  approximans 

Along  Poisson  brook;  along  Choc  River;  near  Vieu.x  Fort; 

at  Union  Agricultural  Station. 
Subulinidae 
Subulina  octona 

Castries;  Bois  d'Orange;  near  Choc  River;  at  Monchy;  near 

Union  Agricultural  Station. 
Streptaxidae 
Guletla  bicolor 

Castries  and  vicinity. 
Bulimulidae 
Protoglyptus  aanctaeluciae 

Castries  and  vicinity. 
Bulimulus  guadalupensis 

Castries  and  vicinity;  several  localities  along  the  coast  and 

foothills;  on  banana  trees  and  shrubs  in  Ferrand  Estate. 
Urocoptidae 
Brachypodella  tatei 

Several  localities  along  coast  and  foothills 
Camaenidae 
Pleurodonte  orbiculata 

Castries;   Choc   River   area;   several   localities   between 

Castries  and  northern  tip  of  island;  Bois  d'Orange;  Cul  de 

Sac  Valley;  Dennery. 
Veronicellidae 
Vaginulus  occidental  is 

Cul  de  Sac  Valley  on  banana  trees;  Ferrand  Estate  on 

banana   trees;    hills   overlooking   Bois   d'Orange;   along 

Poisson  Brook. 
Veroni.cella  floridana 

Several  localities  in  the  northern  half  of  island,  sometimes 

with  V.  occidentalis 

cylidae,  Ampullaridae  and  Hydrobiidae  by  one 
species  each.  Of  the  terrestrial  snails  the  Suc- 
eineidae and  Bulimulidae  were  represented  by  2 
species  each,  while  the  Helicinidae,  Subulinidae, 
Streptaxidae,  Urocoptidae  and  Camaenidae 
were  represented  by  one  species  each.  Two 
species  of  slugs  found  belonged  to  the  family 
Veronicellidae. 

The  planorbid  Biomphalaria  glabrata  (Say, 
1818),  intermediate  host  of  Schistosoma  man- 
soni,  was  found  in  20  localities.  Specimens, 
which  are  up  to  about  10  mm  in  diameter,  never 
attain  the  large  size  of  those  found  in  Puerto 
Rico,  the  Dominican  Republic  or  South  America. 
Accordingly,  F.  C.  Baker  (1945)  showed  figures 


of  B.  glabrata  from  Saint  Lucia  and  commented 
that  they  were  immature.  Examination  of  shell 
features  and  anatomical  details  of  biompha- 
larids  from  various  drainage  systems  indicated 
that  B.  glabrata  is  the  only  species  of  the  genus 
Biomphalaria  that  exists  on  Saint  Lucia.  It  is 
known  that  species  such  as  B.  havanensis.  B. 
helophila  and  B.  straminea  occur  on  some  of  the 
Lesser  and  Greater  Antilles,  alone  or  in  addition 
to  B.  glabrata.  The  snail  was  not  found  in  fast- 
flowing  waters,  but  frequents  and  breeds  on 
flooded  river  and  ravine  (brook)  embankments, 
in  swampy  areas  close  to  rivers,  in  side  pools 
and  backwaters  of  rivers  and  ravines  protected 
by  rocks  and  vegetation,  in  drainage  ditches  of 
banana  fields,  and  in  concrete-lined  channels. 
Sizable  colonies  were  always  found  upstream 
rather  than  near  the  mouth  of  a  river. 

The  planorbids  Drepanotrema  depressissi- 
mum.  (Moricand,  1839)  and  D.  lucidum  (Pfeiffer, 
1839)  live  in  ponds  rich  in  aquatic  vegetation. 
The  lymnaeid  Fossaria  cubensis  (Pfeiffer,  1839) 
is  the  snail  host  of  Fasciola  hepatica  which  is 
common  on  the  island.  Wet  mud  outside  the 
water,  with  or  without  vegetation,  represents 
the  typical  habitat  of  this  lymnaeid,  although  it 
was  also  found  in  narrow  drainage  ditches  in 
banana  fields,  in  shallow  water,  and  in  narrow 
concrete-lined  channels. 

The  Saint  Lucian  material  of  the  hydrobiid 
Pyrgophorus  parvulus  (Guilding,  1828)  exhibits 
variations  common  among  individuals  of  this 
species.  Some  specimens  have  evenly  rounded 
and  smooth  whorls,  whereas  other  specimens 
have  prominent  spiral  carinae  situated  a  short 
distance  between  the  sutures,  and  each  carina 
carries  a  number  of  pronounced  spines. 

The  land  snails  and  slugs  found  during  the 
survey  seemed  to  be  widely  distributed, 
especially  in  the  island's  northern  two-thirds 
where  the  tropical  vegetation  and  coastal 
deposits  of  limestone  exist.  Species  such  as 
Subulina  octona  (Bruguiere,  1789),  Pleurodonte 
orbiculata  (Ferussac,  1821),  Bulimulus 
guadaloupensis  Bruguiere,  1792  and  Protoglyp- 
tus  sanctaeluciae  (E.A.  Smith,  1889)  occur  in 
abundance  in  the  most  populated  town, 
Castries.  The  several  hills  on  which  the  town  is 
located  and  the  coastal  strip  harbor  large  col- 
onies of  these  snails.  Banana  plantations  in 
several  parts  of  the  island  support  large  popula- 


146    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


tions  of  helicinids,  bulimulids,  and  veronicellid 
slugs. 

Discussion 

The  distribution  of  the  freshwater  snails  on 
Saint  Lucia  seems  to  be  related  to  characteris- 
tics of  the  habitat,  such  as  the  stream  gradient, 
water  velocity,  rainfall,  and  salinity.  For  exam- 
ple, B.  glabrata  was  not  found  in  parts  of 
streams  with  steep  gradients  or  in  streams  close 
to  the  sea.  The  nearest  colony  was  found  at 
about  400  yards  from  the  coast.  The  bionomics 
of  S.  glabrata  on  Saint  Lucia  were  investigated 
by  Sturrock  (1973,  1974).  Neritina  spp., 
however,  seem  to  withstand  higher  salinity  than 
the  other  freshwater  snails.  The  drepanotre- 
matids  were  found  to  be  almost  exclusively 
pond-inhabiting  species,  the  exception  being  the 
occurrence  of  Drepanotrenia  depressissimum 
once  in  a  drainage  ditch  and  once  in  a  slow- 
flowing  brook. 

The  distribution  of  the  land  snails  on  this  and 
other  Antillean  islands  follows  the  usual  correla- 
tion between  the  distribution  of  species  and  the 
geological  formation  of  the  area  in  which  they 
occur.  Species  belonging  to  Brachypodella, 
Gulella,  and  Subulina  are  restricted  to  the 
limestone  coastal  land  and  adjacent  hills. 
Pleurodonte  orbiculata  (Ferussac,  1821), 
Bulimulus  guadaloupensis  (Bruguiere,  1792), 
and  Protoglyptus  sanctaeluciae  (E.A.  Smith, 
1889)  also  inhabit  these  areas,  but  they  are  addi- 
tionally found  in  the  igneous  mountains  in  the 
highlands. 

The  main  species  of  freshwater  and  terrestrial 
snails  on  Saint  Lucia  also  occur  on  certain  other 
Caribbean  islands.  The  faunal  relationship  of 
Saint  Lucia  to  other  Caribbean  islands  and  to 
the  mainland  in  Central  and  South  America 
deserves  comment.  Several  theories  have  been 
postulated  to  explain  the  faunal  and  floral 
similarities  among  the  islands  of  the  West  Indies 
on  the  one  hand  and  among  those  of  the  islands 
and  the  fauna  and  flora  of  Central  and  South 
America  on  the  other  hand.  Among  the  theories 
postulated  is  the  past  existence  of  land  l)ridges 
(Ihering,  1931),  or  isthmian  links  (Schuchert, 
1935)  between  the  islands  themselves  and  be- 
tween the  islands  and  the  mainland  of  Central 
and  South  America.  Snails  have  played  an  im- 
portant part  in  helping  to  correlate  the  geo- 


logical history  and  the  zoogeography  of  the 
islands.  Certain  species,  however,  such  as  the 
cosmopolitan  Subulina  spp.  and  others  are  of  lit- 
tle value  in  providing  evidence  of  previous  land 
connections.  The  land  operculate  snails  offer 
evidence  of  zonal  grouping  within  the  West 
Indies.  Surveys  on  Saint  Lucia  revealed  the 
presence  of  only  one  land  operculate,  Helicina 
fasciata  Lamarck,  1818-1822.  Thompson  (1967) 
described  another  operculate  from  Saint  Lucia, 
Lifhacaspis  xanthoglauca  (family  Cyclophori- 
dae).  In  general,  however,  there  is  a  scarcity  of 
land  operculate  snails  on  the  Lesser  Antillean 
islands  and  South  America  as  compared  to  a 
preponderance  of  land  operculates  on  the 
Greater  Antilles  islands. 

Among  the  freshwater  snails,  the  planorbids 
show  relationships  with  the  South  American 
fauna.  Drepanotrem.a  depressissimum  (Mori- 
cand,  1839)  is  typically  South  American.  B. 
glabrata  is  found  on  some  islands  of  both  the 
Lesser  and  the  Greater  Antilles,  and  its  range 
extends  into  South  America.  In  the  Antilles,  B. 
glabrata  occurs  on  Saint  Lucia,  Martinique, 
Guadeloupe,  St.  Kitts,  Antigua,  Dominica, 
Hispaniola,  and  Puerto  Rico.  Ampullaria  glauca 
(Linnaeus,  1758)  is  also  South  American. 

The  melaniid  Thiara  granifera  (Lamarck, 
1822)  was  introduced  by  Prentice  (1983)  in  field 
trials  in  marshes  and  streams  for  experiments 
on  biological  control  oiB.  glabrata.  Whether  T. 
granifera  still  exists  in  these  habitats  or  has 
spread  to  other  habitats  in  Saint  Lucia  is  not 
known.  Jordan  (1985)  listed  the  sphaeriid 
l)ivalve  Pisidium  punctiferum  (Guppy)  as  a 
member  of  the  freshwater  fauna  of  the  island, 
but  it  was  not  encountered  in  my  surveys.  How- 
ever, his  list  did  not  include  the  terrestrial  oper- 
culate Helicina  fasciata  Lamarck,  nor  the  slug 
Veronicella  floridana  (Leidy)  which  were  both 
common  in  the  material  that  I  collected. 

LITERATURE  CITED 

Baker,  F.  C.  1945.  TheMolluncan  Family  Planorhidar.  I'ni- 
versity  of  Illinois  Press,  Urbana,  Illinois. 

Gomez,  J.  D.,  Vargas,  M.,  and  Malek.  E,  A.  1986.  Fresh- 
water mollusks  of  the  Dominican  Republic.  The  Niiittilus 
100  (in  press). 

(Uiyani,  A.  and  Pointier,  J.  P.  1979.  Faune  malacolofjique 
diiicaquicole  et  vecteurs  de  la  schistosome  intestinale  en 
Martinique  (Antilles  Francaises).  Ann.  Pnra.'^itol.  (Paris): 
54:  193-205. 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    147 


Ihering,  H.  von.  1931.  Land  bridges  across  the  Atlantic  and 
Pacific  Oceans  during  the  Kainozoic  era.  Quart.  J.  Geol. 
Soc.  London  87:376-391. 

Jordan,  P.  1985.  Schistosomiasis.  The  St.  Lucia  Project. 
Cambridge  University  Press,  Cambridge. 

Malel<.  E.  A.  1962.  Report  on  precontrol  studies  of  bil- 
harziasis  in  St.  Lucia.  Unpublished  Document,  Pan 
American  Health  Organization,  Washington,  D.C.:  1-18. 

1963.  Report  on  a  precontrol  survey  of  bil- 

harziasis  in  a  pilot  study  area  of  St.  Lucia.  Second  visit. 
Unpublished  Document,  AMRO-1.55,  Pan  American 
Health  Organization,  Washington,  D.C.:  1-9. 

.  1965.  Freshwater  and  land  snails  of  St.  Lucia, 


the  West  Indies.  Amer.  Malacot.  Union.  Annual  Report. 
p.  38. 
1980.  Snail-Tra7isntitted  Parasitic  Diseases. 


Vol.  I.  CRC  Press,  Boca  Raton,  Florida. 
Pointier,  J.  P.  1974.  Faune  malacologique  dulcaquicole  de 

I'ile  de  la  Guadeloupe  (Antilles  Francaises).  Bull.  Mus. 

Hist.  Natur..  Paris,  3rd  Ser.,  No.  235,  Zool.,  159:905-933. 
Prentice,    M.    A.    1983.    Displacement    of   Biomphalaria 

glabrata  by  the  snail  Thiara  granifera  in  field  habitats  in 

St.    Lucia,    West    Indies.    Ann.    Trap.    Med.    Parasitol. 

77:51-59. 
Robart,  G.,  Mandahl-Barth,  G.  and  Ripert,  C.  1976.  Inven- 

taire,  repartition  geographique  et  ecologie  des  mollusques 

dulcaquicoles  d'Haiti  (Carabes).  Hnliotts  8:159-171. 


Schuchert,  C.  1935.  Historical  Geology  of  the  Antillean- 
Caribbean  Region.  John  Wiley  &  Sons,  Inc.,  New  York. 

Smith,  E.  A.  1889.  On  the  Mollusca  collected  by  Mr.  G.  A. 
Ramage  in  the  Lesser  Antiles.  Report  III.  Ann.  Mag. 
Natur.  Hist.  (6th  Ser.)  3:400-405. 

1895.  Report  on  the  land  and  freshwater  shells 

collected  by  Mr.  H.  H.  Smith  at  St.  Vincent,  Grenada  and 
other  neighboring  islands.  Proc.  Malacol.  Soc.  London 
1:300-322. 

Smith,  E.  A.  and  Feilden,  H.  W.  1891.  A  list  of  the  land  and 
freshwater  shells  of  Barbados.  Ann.  Mag.  Natur.  Hist. 
(6th  Ser.)  5:247-257. 

Starmuhlner,  F.  1985.  Erstfunde  von  drei  Arten  von  Suss- 
wasser  Gastropoden  auf  den  Inseln  Guadeloupe,  Dominica 
und  Martinique  (Kleine  Antillen).  Heldia  1:55-58. 

Sturrock,  R.  F.  1973.  Field  studies  on  the  transmission  of 
Schistosoma  mansoni  and  on  the  bionomics  of  its  inter- 
mediate host,  Biomphalaria  glabrata,  on  St.  Lucia,  West 
Indies.  Int.  Jour.  Parasitol.  3:175-194. 

.  1974.  Ecological  notes  on  habitats  of  the  fresh- 
water snail  Biomphalaria  glabrata,  intermediate  host  of 
Schistosoma  mansoni,  on  St.  Lucia,  West  Indies.  Carib- 
bean Jour.  Sci.  14:149-161. 

Thompson,  F.  G.  1967.  A  new  cyclophorid  land  snail  from 
the  West  Indies  (Prosobranchia)  and  the  discussion  of  a 
new  subfamily.  Proc.  Biol.  Soc.  Washington  80:13-18. 


ON  THE  REDISCOVERY  OF  TERAMACHIA  MIRABILIS 

(CLENCH  AND  AGUAYO,  1941),  AND  ITS 
RELATIONSHIP  TO  OTHER  CALLIOTECTINE  VOLUTES 

William  K.  Emerson  and  Walter  E.  Sage  III 

Department  of  Invertebrates 

American  Museum  of  Natural  History 

New  York,  N.  Y.  10024 


ABSTRACT 
A  second  known  specimen  o/Teramachia  mirabilis  (Cleyich  and  Aguayo,  19Jfl)  is 
here  recorded  from,  deep  water  (UGS.U  meters)  on  the  Little  Bahama  Bank.  Com- 
parisons of  this  dead-collected,  crabbed  speciynen  are  jnade  with  illustrations  and 
descriptions  of  the  holotype,  which  was  live-taken  in  521.3  meters  off  Matayizas, 
Cuba,  in  1939.  Largely  on  the  similarity  of  shell  characteristics,  this  New  World 
species  is  retained  in  the  genus  Teramachia,  as  is  Calliotectum  fischeri  Olsson, 
1964,  from  the  Pliocene  of  Ecuador.  Examination  of  the  typological  specimens  of 
Calliotectum  vernicosum  Dall,  1890,  for  which  a  lectotype  is  here  designated,  con- 
firms the  distinction  between  Calliotectum  Dall,  1890  and  Teramachia  Kuroda, 
1931. 

Zoogeographically,  Teramachia  is  confined  mostly  to  the  western  Pacific  and  is 
known  to  date  froyn  the  Neogene  of  Japan  as  ivell  as  Ecuador.  Thus  the  widely 


148    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


separated  ynodem  distributional  pattern  of  the  genus  Teramachia  (six  species  in 
the  western  Pacific,  one  in  the  western  A  tlantic)  suggests  that  T.  mirabilis  is  a  sur- 
viving element  of  a  presently  disjunct  Pacific  fauna. 


Howellia  mirabilis  Clench  and  Aguayo,  (1941, 
pp.  177-178,  pi.  14,  fig.  2;  Weaver  and  duPont, 
1970,  p.  178,  pi.  76  A,  B)  was  described  on  the 
basis  of  a  single,  live-collected  specimen 
dredged  at  Atlantis  Station  No.  3483,  off 
Matanzas.  Matanzas  Province,  Cuba  (23°12'N, 
81°23'W),  in  521.3  meters.  The  soft  parts  unfor- 
tunately were  not  recovered  in  a  condition  that 
could  be  preserved,  and  the  shell  was  cleaned 
and  dried  during  the  trip.  Regrettably,  the 
holotype  (catalog  #135291)  cannot  be  located  in 
the  collection  of  the  Museum  of  Comparative 
Zoology  and  is  presumed  lost  (teste  D.  Backus). 
This  taxon  was  proposed  as  the  type  species  for 
the  monotypic  genus  Howellia  by  Clench  and 
Aguayo  (1941,  p.  177).  Howellia  was  placed  pro- 
visionally by  Clench  and  Aguayo  (op.  cit.)  in  the 
family  Fasciolariidae,  in  the  absence  of  know- 
ledge of  the  radula  and  soft-part  anatomy.  Sub- 
sequently, Clench  and  Turner  (1964,  p.  177) 
assigned  Howellia  to  the  Volutidae,  subfamily 
Calliotectinae,  following  the  placement  of 
Pilsbry  and  Olsson  (1954,  p.  19),  based  on  shell 
characters.  Clench  and  Turner  (1964,  p.  178), 
however,  noted  the  resemblance  in  shell  mor- 
phology of  Howellia  mirabilis  to  the  western 
Pacific  genus  Teramachia  Kuroda,  1931.  Later 
Weaver  and  duPont  (1970,  p.  176)  placed 
Howellia,  1941,  in  the  synonymy  of  Teramachia, 
1931,  where  it  was  retained  by  Emerson  (1985, 
pp.  102,  103). 

Recently  Harry  G.  Lee  of  Jacksonville, 
Florida  kindly  called  our  attention  to  a  crabbed 
specimen  of  Teramachia  mirabilis  in  the  collec- 
tion of  the  Indian  River  Coastal  Zone  Museum 
(IRCZM)  at  Harbor  Branch  Oceanographic  In- 
stitution, Inc.,  Fort  Pierce,  Florida.  This 
specimen,  apparently  only  the  second  known  for 
this  taxon,  was  found  in  a  sediment  sample  col- 
lected on  a  Johnson-Sea-Link  I  submersible  dive 
during  R/V  Johnson  Cruise  159,  on  October  4, 
1983.  The  bucket  had  been  left  on  the  bottom  for 
408  days  to  collect  accumulating  sediment  near 
Black  Rock,  Little  Bahama  Bank  (26°16.5'N, 
77°38.5'W)  in  465.4  meters,  by  C.  M.  Hoskin 
and  J.  K.  Reed.  The  specimen  presumably  was 
carried  into  the  20-liter  plastic  bucket  by  the 


hermit  crab  occupying  the  shell. 

Although  long-dead  and  now  discolored  from 
the  muddy  sediment,  this  specimen,  IRCZM 
#065:02138  (figs.  5,  6),  compares  favorably  with 
the  description  and  illustration  of  the  holotype. 
The  present  specimen  is  larger  (125  vs.  93  mm 
in  height)  and  has  weaker  axial  sculpture  on  the 
body  whorl.  In  outline  and  in  the  development  of 
axial  sculpture  on  the  body  whorl  Teranfiachia 
mirabilis  most  closely  resembles  the  western 
Pacific  Teramachia  johnsoni  (Bartsch,  1942,  p. 
12,  pi.  2,  fig.  3;  holotype  illustrated  in  color  by 
Weaver  and  duPont,  1970,  pi.  750,  H;  cf.  figs.  5, 
6  with  figs.  7,  8  herein).  Teramachia  johnsoni, 
however,  has  a  tannish  brown  shell  with  a 
darker  aperture,  whereas  the  holotype  of  Tera- 
machia mirabilis  is  whitish  and  tinged  with  a 
faint  brownish  red.  The  aperture  is  light  brown- 
ish red  within. 

The  discovery  of  the  newly  recognized  speci- 
men of  Teramachia  mirabilis  extends  the 
known  provenance  of  this  species  northwest- 
ward from  the  vicinity  of  Cay  Sal  Bank,  off 
northern  Cuba,  to  the  Little  Bahama  Bank. 
Specimens  can  be  expected  to  occur  at  suitable 
depths  elsewhere  on  these  and  other  regional 
banks. 

In  the  absence  of  anatomical  data  on  the  type 
species  of  Howellia  it  seems  prudent  to  refer 
this  New  World  species  to  the  genus  Tera- 
machia on  the  basis  of  shell  characters  held  in 
common  with  the  western  Pacific  species  of  this 
group  of  calliotectine  volutes  (Weaver  and 
duPont,  1970,  p.  177,  Emerson,  1985,  p.  103).  A 
more  precise  systematic  assessment  oi  Howellia 
must  await  knowledge  of  the  soft  parts  and 
radular  characters  of  Teraynachia  mirabilis. 

Clench  and  Turner  (1964,  p.  178)  briefly  noted 
the  close  resemblance  of  the  shell  morphology  of 
Howellia  mirabilis  to  that  of  the  Japanese 
species  of  Teramachia,  and  they  stated,  "Rela- 
tionships of  this  sort  from  widely  separated 
areas  are  unusual,  but  they  do  exist".  Shortly 
thereafter,  Olsson  (1964,  p'.  129,  pi.  23,  fig.  4) 
described  Calliotectum  fischeri  from  late 
Pliocene  rocks  of  the  Esmeraldas  formation  of 
Ecuador.  The  holotype  is  based  on  a  52.8  mm 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    149 


150    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


fragmental  specimen,  which  was  estimated  to  be 
"between  six  and  eight  inches  in  length"  before 
the  badly  fractured  specimen  was  extracted 
from  the  matrix.  Olsson  compared  his  specimen 
to  Prodallia  dalli  Bartsch,  1942  from  the  Philip- 
pines and  he  referred  Teramachia  Kuroda,  1931 
and  Prodallia  Bartsch,  1942  to  the  synonymy  of 
Calliotectum  Dall,  1890.  We  have  examined  the 
holotype  and  six  fragmental  specimens  of 
Olsson's  "Calliotectum"  fischeri  from  near  the 
type  locality  (Tulane  Geol.  Dept.  localities  1397 
[figs.  3,  4,  herein]  and  1402,  and  W.  D.  Pitt  Coll. 
locality  9A-B3-915,  all  from  the  Esmeraldas  for- 
mation). These  specimens  are  from  a  deep-water 
facies  and  confirm  the  allocation  of  Olsson's 
taxon  to  the  genus  Teramachia.  The  only  other 
known  extinct  species  of  Teramachia  was 
described  from  the  Mio-Pliocene  [?Pliocene] 
Shimajiri  formation  of  Okinawa,  Japan  as  T. 
shinzatoensis  (MacNeil,  1961,  p.  96,  pi.  9,  fig.  1; 
Rehder,  1972,  p.  9,  figs.  5,  6).  It  was  compared 
by  the  describer  and  Dr.  Rehder  with  T. 
johnsoni  (Bartsch,  1942). 

The  placement  by  Olsson  (1964,  p.  128)  of  his 
Pliocene  Ecuadorian  fossil  (Teramachia 
fischeri)  in  the  monotypic  genus  Calliotectum 
Dall  (1890,  p.  304)  requires  comment.  Dall 
(1890,  p.  305,  pi.  5,  fig.  8)  considered  the  type 
species  of  Calliotectum  (C.  vernicosmn  Dall, 
1890)  to  be  pleurotomoid,  but  he  noted  the  shell 
lacked  an  anal  notch  and  a  fasciole.  Subsequent- 
ly, Pilsbry  and  Olsson  (1954,  p.  19,  fig.  16,  rachi- 
dian  tooth)  proposed  the  volutid  subfamily 
Calliotectinae  with  Calliotectum  the  type  genus, 
on  the  basis  of  shell  and  radular  characters,  the 
radular  ribbon  being  uniserial  with  a  tricuspid 
rachidian  tooth.  Pilsbry  and  Olsson  (1954,  op. 
cit.)  also  referred  Teramachia  Kuroda,  1931, 
with  Prodallia  Bartsch,  1942  as  junior 
synonym,  to  Calliotectinae.  Weaver  and  duPont 
(1970,  pp.  175,  176)  added  Howellia  Clench  and 
Aguayo,  1941  to  the  synonymy  of  Teramachia, 
1931. 

Dall  (1890,  p.  305)  based  Calliotectum  ver- 
nicosum  on  five  specimens  dredged  by  the 
"Albatross",  in  1888.  Three  of  these  were  taken 


at  station  2807,  in  1485  meters  from  the  type 
locality  near  the  Galapagos  Islands  (00°24'00"S, 
89°06'00"W).  From  this  lot  are  Dall's  figured 
specimen  USNM  #96555  (1890,  pi.  5,  fig.  8; 
Abbott  and  Dance,  1982,  p.  224)  here  selected  as 
lectotype  (figs.  9,  10),  and  two  paralectotypes: 
USNM  #633904,  and  DMNH  #10135  (Weaver 
and  duPont,  1970,  pi.  75,  A,  B)  -  both  ex  USNM 
#96555.  The  two  remaining  paralectotypes 
(USNM  #97068)  were  dredged  at  station  2793, 
in  1355  meters  off  the  coast  of  Ecuador 
(01°03'00"N,  18°15'00"W). 

These  typological  specimens  are  very  thin- 
shelled,  with  inflated  whorls,  the  largest 
specimen  being  the  lectotype  with  a  height  of 
47.5  mm  (spire  incomplete)  and  with  6V2  post- 
nuclear  whorls,  and  the  smallest  paralectotype 
being  24.8  mm  in  height  (spire  incomplete)  with 
5  postnuclear  whorls.  Although  superficially 
resembling  the  shells  of  Teramachia.  specimens 
of  Calliotectum.  of  the  same  stage  of  growth 


FIGS.  9,  10.  Calliotectum  reniira^um  Dall.  1890,  near  the 
GalapaKos  Islands,  in  1485  meters,  lectotype  USNM  #96555; 
X    1. 


FIGS.  1,2.  T.  dalli  daydoni  Poppe,  1986,  off  Port  Hedland,  northwestern  Australia,  in  450  meters,  AMNH  #221239.  FIGS.  3, 
4.  Teramachia  fifickeri  (Olsson,  1964),  Quebrada  Camerones,  Province  of  Esmeraldas,  Ecuador,  Tulane  Geol.  Dept.  loc.  1397, 
Pliocene  (early  whorls  and  basal  whorls  missing).  FIGS.  5,  6.  T.  mirabilis  (Clench  and  Aguayo,  1941),  Little  Bahama  Bank,  in 
465  meters,  IRCZM  #065:02138.  FIGS.  7,  8.  T.  johnsoni  (Bartsch,  1942),  off  Panglao,  Bohol.  Philippines,  in  365  meters, 
AMNH  #219986.  FIGS.  1-8,  approximately  x  1. 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    151 


have  expanded,  bulbous  whorls  and  appear  to 
represent  a  separate  calliotectine  lineage  (cf. 
figs.  9,  10  with  figs  1,  2  and  figs.  3,  4).  Therefore 
we  reject  Olsson's  placement  of  Teramachia 
Kuroda,  1931  and  Prodallia  Bartsch,  1942  in 
the  synonymy  of  Calliotectum  Dall,  1890. 

The  widely  disjunct  modern  distributional  pat- 
tern of  Teramachia,  six  species  in  the  western 
Pacific  and  one  in  the  western  Atlantic  (Emer- 
son, 1985;  Bouchet,  1986;  Poppe,  1986),  with  ex- 
tinct species  in  the  Mio-Pliocene  of  Okinawa  and 
the  Pliocene  of  Ecuador,  presents  some  prob- 
lems in  zoogeographic  interpretation.  The  dis- 
tribution of  these  deep-water  mollusks  is  poorly 
known  and  the  genus  may  be  of  wider  occur- 
rence than  the  available  data  suggest.  Within 
the  past  few  months,  Teramachia  dalli  claydoni 
Poppe  (1986)  was  described  from  the  continen- 
tal slope  of  northwestern  Australia  and  an  ap- 
parently new  species  of  Teramachia  inhabiting 
deep  water  off  New  Caledonia  was  reported 
(Bouchet,  1986).  Additional  species  may  even- 
tually be  found  living  in  the  eastern  Pacific  and 
elsewhere  in  the  Atlantic  Basin.  The  presence  of 
Teramachia  mirabilis  living  in  the  western 
Atlantic,  however,  may  be  an  example  of  a 
Pacific  faunal  element  which  survived  after 
being  carried  into  the  Caribbean  region  on  the 
East-Pacific-Caribbean  plate  to  its  present  posi- 
tion adjacent  to  Cuba  (see  Durham,  1985).  Ac- 
cording to  Sykes  et  al.  (1982),  the  Caribbean 
plate  has  moved  east-northeast  about  1,400  km 
since  late  Eocene  time.  Perhaps  the  precursors 
of  T.  mirabilis  were  carried  into  the  western 
Atlantic  on  this  plate.  Durham  (1985)  cites  addi- 
tional examples  of  Pacific  faunal  elements  that 
apparently  moved  with  this  plate  into  the  Carib- 
bean area. 

Acknowledgments 

In  addition  to  Dr.  Harry  Lee,  we  thank  Paula 
Mikkelsen,  Indian  River  Coastal  Zone  Museum, 
Harbor  Branch  Oceanographic  Institution,  Inc., 
Fort  Pierce,  Florida,  William  D.  Pitt,  Sacra- 
mento, California,  Drs.  Emily  and  Harold 
Yokes,  Tulane  University,  and  Drs.  Richard  S. 
Houbrick  and  Thomas  D.  Waller,  National 
Museum  of  Natural  History,  Smithsonian  In- 
stitution, for  the  loan  of  specimens  in  their  care. 
We  also  thank  Drs.  Joel  Cracraft,  Leslie  F. 
Marcus,  Malcolm  McKenna,  and  Richard  H. 
Tedford  for  an  exchange  of  views  on  the  bio- 


geographical   significance   of  Caribbean   plate 
tectonics. 

We  are  indebted  to  our  colleagues  in  the  De- 
partment of  Invertebrates,  American  Museum 
of  Natural  History  for  their  contributions  to  this 
study:  Stephen  M.  Butler  for  the  photography 
and  Stephanie  Crooms  for  word-processing  the 
manuscript. 

LITERATURE  CITED 

Abbott,  R.  T.  and  Dance,  S.  P.  1982.  Compendium,  of  Sea- 
shells.  New  York,  411  pp.,  illus.  in  color. 

Bartsch,  P.  1942.  Some  deep-sea  Philippine  volutids.  The 
Nautilus  56(1):9-13,  pi.  2. 

Bouchet.  P.  1986.  Oceanographic  campaigns  in  New  Cale- 
donia. Rossiniana,  Bull.  L'Assoc.  Conchyl.  De  Nouvelle- 
Caledonie  31;3-8,  illus. 

Clench,  W.  J.  and  Aguayo,  C.  G.  1941.  Notes  and  descrip- 
tions of  new  deep-water  Mollusca  obtained  by  the 
Harvard-Havana  Expedition  off  the  coast  of  Cuba.  IV. 
Mem.  Soc.  Cubana  Hist.  Nat.  15(2):177-180,  pi.  14, 

Clench,  W.  J.  and  Turner,  R.  D.  1964.  The  subfamilies  Volu- 
tinae,  Zidoninae,  Odontocymbiolinae,  and  Calliotectinae  in 
the  western  Atlantic.  Johnsonia  4(43):129-180,  pis. 
80-114. 

Dall,  W.  H.  1890  [1889].  Scientific  results  of  explorations  by 
U.  S.  Fish  Commission  Steamer  Albatross.  VII.  Prelimi- 
nary report  on  the  collection  of  Mollusca  and  Brachiopoda 
obtained  in  1887-'88.  Proc.  U.  S.  Nat.  Mu.s.  12(737): 
219-362.  pis.  5-14. 

Durham,  J.  W.  1985.  Movement  of  the  Caribbean  plate  and 
its  importance  for  biogeography  in  the  Caribbean.  Geology 
13(2):123-125,  2  figs. 

Emerson,  William  K.  1985.  Teramachia  dupreyae  new 
species,  from  off  Western  Australia  (Gastropoda:  Voluti- 
dae).  The  Nautilus  99(4):102-107,  8  figs. 

Kuroda,  T.  1931.  Two  new  species  of  Volutacea.  Venus  3(1): 
45-49,  3  figs. 

Olsson,  A.  A.  1964.  Neogene  mollusks  from  northwestern 
Ecuador.  Paleont.  Res.  Inst.,  256  pp.  36  pis, 

MacNeil,  F.  S.  1961.  Tertiary  and  Quaternary  gastropods  of 
Okinawa.  U.  S.  Geol.  Surv.  Prof.  Paper  339  ("I960"):  i-iv 
+  1-148,  pis.  1-19  (distributed  March  17,  1961). 

Pilsbry,  H.  A.  and  Olsson,  A.  A.  1954.  Systems  of  the 
Volutidae.  Bull.  Amer.  Paleont.  35(152):271-306,  pis. 
2.5-28. 

Poppe,  G.  1986.  A  novel  species  and  a  novel  subspecies  of 
Volutidae  from  northwestern  Australia.  .4 pfx.  Inform.  Sci. 
Soc.  Beige  de  Malac.  l(l):27-36,  pis.  1-3. 

Rehder.  H.  A.  1972.  Some  notes  on  the  genus  Teramachia 
Volutidae:  Calliotectinae).  The  Veliger  15(1):7-10,  figs. 
1-7. 

Sykes,  L.  R.,  McCann,  W.  R.,  and  Kafka,  A.  L.  1982.  Motion 
of  Caribbean  plate  during  last  7  million  years  and  implica- 
tions for  earlier  Cenozoic  movements.  Jour.  Geophysical 
Res.  87(B13):106.56-10676. 

Weaver.  C.  S.  and  duPont,  J.  E.  1970.  Living  Volutes:  a 
monograph  of  Recent  Volutidae  of  the  world.  Delaware 
Mus.  Nat.  Hist.  Mong.  Ser.,  No.  1,  xv  +  375  pp.,  79  col. 
pis.,  44  figs.,  13  maps. 


152    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


REMINISCENCES 

[About  a  hundred  years  ago  The  Nautilus  published  many  anecdotal  accounts, 
but  as  the  journal  became  more  technical  there  was  rarely  an  opportunity  to  add  a 
personal  touch.  Because  this  100th  anniversary  volume  is  dedicated  to  mala- 
cologists  who  have  contributed  in  the  past,  we  are  including  a  few  personal 
reminiscences  by  friends  of  William  J.  Clench  and  Joseph  Rosewater.  To  the  list 
of  departed  fellow-malacologists  we  regretably  must  add  the  name  of  that 
distinguished  freshwater  expert,  Henry  van  der  Schalie,  whose  touching  obituary 
is  included  in  this  number  of  The  Nautilus. -R.T. A.,  editor]. 


To  Joe  Rosewater— from  R.  Tucker  Abbott 

[read  at  the  1985  memorial  tribute  at  the 
Smithsonian  Institution] 
The  corridors  of  malacology  may  be  darkened 
today  with  the  passing  of  our  colleague  and 
friend,  Joe  Rosewater,  but  we  and  generations 
to  follow  will  find  many  bright  rooms  off  to  the 
side  that  are  filled  with  Joe's  warm  kindness  and 
illuminating  scientific  labors. 

Calliostoma  rosewateri- one  of  the  most  beau- 
tiful seashells  of  the  Caribbean  -  was  appropri- 
ately  named   after   Joe   by   Drs.   Clench   and 
Turner  twenty  five  years  ago.  Joe  loved  and 
understood  the  human  qualities  of  his  colleagues 
and  a  year  later,  in  his  famous  biography  of 
Calvin  Goodrich,  he  quoted  two  letters  written 
to  his  mentor,  Bill  Clench: 
"Thank  you  for  naming  the  new  species  for 
me.  In  the  days  to  come  when  the  new  genera- 
tion rolls  up  its  sleeves  to  undo  the  work  of 
this  one,  as  this  one  is  undoing  that  of  [tax- 
onomic]  workers  who  thought  they  had  done  a 
good,  decent  job,  some  one  will  note  the  name 
and  speculate  a  bit  about  the  guy  it  honors, 
and  that  will  be  in  the  nature  of  fame." 
And  Joe,  with  a  sense  of  humor,  went  on  to 
quote  another  prophetic  letter: 
"Your  students  [meaning  Clench's  students] 
are  of  a  high  order.  I  was  delighted  with  them, 
and  would  steal  them  away  if  I  knew  just  how 
to  go  about  it.  After  all,  you've  more  students 
than  you  should  have,  considering  the  rarity  of 
boys  interested  in  mollusks.  I  wish  you  would 
tell  them  that  I  consider  it  far  more  of  a 
pleasure    to    have    met    them    and    I    have 
delighted    in    their   friendliness   and    intelli- 
gence,   whatever   the   police    may   think   of 
them." 

Just  a  month  ago  I  received  a  letter  from  a 
now-retired,  U.  S.  National  Museum  paleon- 
tologist living  in  Florida: 


"Joe  really  did  a  fine  job  of  running  the  Divi- 
sion of  Mollusks,  and  one  could  always  write 
to  him  for  help  and  expect  a  prompt  reply.  Joe 
was  a  careful  and  meticulous  researcher  as 
you  well  know." 

Joseph  Rosewater  was  born  in  Claremont, 
New  Hampshire,  on  September  18,  1928.  He 
was  of  the  same  New  England  breed  of  mala- 
cologists  as  the  Binneys,  Dall,  Morse,  C.  B. 
Adams,  A.  E.  Verrill  and  W.  J.  Clench.  Follow- 
ing his  early  college  days  at  the  University  of 
New  Hampshire,  Joe  began  his  advanced  train- 
ing in  malacology  and  museum  curating  at 
Harvard  University.  He  was  a  favorite  student 
of  Clench's,  and  as  Goodrich  would  say,  was 
unknown  to  the  police. 

His  magnificent  service  to  the  Smithsonian  In- 
stitution began  in  1960.  Joe's  twenty-five  years 
of  malacology  was  a  fulfillment  of  James 
Smithson's  bequest  for  the  "increase  and  diffu- 
sion of  knowledge  among  all  men."  He  served  as 
President  of  the  American  Malacological  Union 
in  1968,  and  as  President  of  the  National  Capitol 
Shell  Club  in  1964.  His  contributions  to  the 
leading  malacological  journals  of  America  are 
well-known  to  his  colleagues.  His  travels  in 
search  of  mollusks  to  the  Marshall  Islands, 
Indonesia,  Australia  and  Panama  were  the  envy 
of  all  devoted  shell  collectors.  No  more  gentle 
soul,  with  such  a  quiet  sense  of  humor,  nor  with 
a  more  natural  ability  to  help  both  professional 
or  amateur  malacologists  could  we  all  have 
asked  for.  Let  us  hope  that  his  successors  will 
match  his  sterling  qualities.  Let's  light  up  the 
halls  of  the  National  Museum  once  again. 

About  Bill  Clench -from  Mary  C.  Kline 

[Mary  Kline,  now  of  Honolulu.,  and  her  late 
husband,  George,  were  for  many  years  ardent 
shell  collectors  who  contributed  many  thousands 
of  specimens,  photographs  and  much  biological 


Vol.  100(4) 


October  31,  1986 


THE  NAUTILUS    153 


information  on  motlusks  for-  the  journals. 
Johnsonia  and  Indo-Pacific  Mollusca.] 

Bill  Clench  was  one  of  my  favorite  "ex- 
tremists"-extremely  kind,  caring  and  loyal;  an 
extremely  good  raconteur.  By  his  enthusiasm 
for  collecting  shells  and  his  insisting  that  the 
neophyte  keep  good  field  notes,  he  directed 
many  beginners  down  the  happy  road  to  ex- 
tremely rewarding  and  meaningful  adventures. 

Our  friendship  with  Bill  started  about  forty 
years  ago  when  George  and  I  first  fell  prey  to 
the  spell  of  the  shell  during  a  visit  to  Sanibel.  On 
that  first  day  of  beach-walking  George  found  a 
"different  looking"  shell.  Local  savants  urged  us 
to  write  to  Dr.  Clench  at  Harvard.  The  shell 
turned  out  to  be  an  almost  unheard  of  albino 
Fasciolaria  distans. 

And  so  began  an  encouraging  flood  of  corre- 
spondence in  response  to  our  new  shelling  trips 
to  the  Florida  Keys  and  Bahamas.  Bill  was  in 
those  days  launching  his  new  journal, 
Johnsonia.  We  had  many  live-collected 
specimens,  plenty  of  notes  and  colored  pictures 
of  animals.  "Drop  everything,  come  to  the  MCZ 
[Museum  of  Comparative  Zoology]  for  a  week- 
end and  bring  the  slides."  We  went,  we  talked, 
we  spent  a  long  time  on  the  steps  of  the  museum 
bing  innoculatead  with  Bill's  enthusiasm.  At  his 
home,  his  wife  Julia,  was  a  marvelous  hostess 
who  had  great  patience  and  was  a  perfect  house 
mother  to  all  and  sundry  who  interested  Bill. 

Bill  was  many  things  but  hardly  the  Ivory 
Tower  type.  For  example:  one  day  when  we  ar- 
rived at  the  museum  lab  he  could  hardly  wait  to 
show  us  his  latest  treasure.  We  knew  that  he 
and  his  associates  had  been  working  for  a  long 
time  on  a  seemingly  endless  project -cleaning 
and  rehabilitating  the  myriad  drawers  in  the 
many  huge  specimen  cabinets.  As  each  drawer 
was  finished  a  small  blue  star  was  affixed  to  the 
front,  indicating  that  it  had  been  completed.  At 
last  the  goal  had  been  reached  and  a  party  was 
given  to  celebrate.  Bill  was  presented  with  a 
beautifully  wrapped  gift  package.  Inside  was  a 
suitably  framed  miniature  of  a  real  pair  of  Long- 
John  bottoms  with  blue  stars  glued  all  over 
them!  How  this  wonderful  man  loved  those 
"Blue  Starred  Drawers!" 

He  begged  us  to  become  interested  in  land 
shells  as  well  as  freshwater  mollusks  for,  he 
said,  "the  human  population  will  grow  and  grow 


and  habitats  will  become  neighborhoods -fresh- 
water shells  will  disappear  for  pollution  is  in- 
evitable but  the  sea  will  always  be  with  us."  He 
lived  to  see  the  land  fills  where  once  there  were 
reefs.  Many  fashionable  resorts  now  have 
replaced  sleepy  lagoons  over  much  of  the 
tropical  world. 

What  did  I  learn  from  Bill? -so  many  things - 
and  here  are  just  a  few.  1 .  A  shell  without  data  is 
just  a  pretty  bauble.  2.  Learn  the  scientific 
names,  beach  names  are  generally  local.  3.  Be 
sure  to  learn  the  dangers  of  the  areas  in  which 
you  are  shelling -and  avoid  them.  4.  Join  the 
A.M.U.  and  your  local  Shell  Club.  5.  In  a  foreign 
country  deal  with  the  proper  authorities  and 
remember,  in  your  own  way  you  represent 
America.  6.  Ask  the  museum  curator  how  you 
can  help -leave  the  scientific  side  to  him.  7. 
Keep  accurate  notes  and  charts -be  a  good 
observer.  8.  Don't  take  all  shells  from  one  col- 
ony-leave the  habitat  as  you  found  it.  9.  And 
lastly,  but  far  from  least,  keep  your  sense  of 
humor. 

Remembering  Bill  Clench  — 
from  Bunny  Baker 

[Mrs.  Horace  B.  Baker,  for  many  years  the 
business  manager  of  The  Nautilus,  was  raised 
in  Boston,  and  has  been  an  ardent  shelter  for  over 
fifty  years.] 

I  first  met  Bill  Clench  in  the  fall  of  1933  when 
I  went  to  work  at  the  Boston  Children's 
Museum.  The  museum's  exhibits  were  mostly  on 
natural  history  topics,  and  though  I  was  the 
financial  secretary  and  only  office  worker,  I 
thought  I  should  know  some  natural  history.  My 
lifelong  friend  and  childhood  chum,  who  also 
worked  in  the  museum,  and  I  had  collected 
shells  along  the  Mass.  coast,  and  so  that  seemed 
the  logical  place  to  start.  I  joined  the  Boston 
Malacological  Club,  and  there  met  Bill  who  was 
to  become  a  large  part  of  my  life. 

In  the  fall  of  1934  my  chum,  her  mother,  sister 
and  I  went  to  Miami  for  our  vacation.  Her  family 
returned  home  by  boat  after  a  few  days,  and  we 
two  girls  drove  across  the  Tamiami  Trail  to 
Sanibel  Island  to  collect  shells.  Bill  had  told  us 
so  much  about  it  that  we  wanted  to  collect  there. 
After  a  week  we  drove  back  to  the  east  coast 
and  met  Bill  and  two  of  his  students  who  had 
been  on  a  collecting  trip  to  Cuba.  We  all  drove 


154    THE  NAUTILUS 


October  31,  1986 


Vol.  100(4) 


down  to  Homestead  and  into  the  Everglades 
where  we  camped  out  for  five  days  and  collected 
Liguus.  Bill  helped  us  identify  the  various  color 
forms  and  taught  us  a  lot  about  collecting  and 
cataloguing  our  catch.  It  was  a  wonderful  ex- 
perience and  of  course  we  all  got  really  ac- 
quainted with  each  other.  We  got  hundreds  of 
shells,  and  those  added  to  my  Sanibel  collection 
and  others  from  around  the  world  gave  me  quite 
a  cabinetfull.  They  are  now  in  Harvard's 
Museum  of  Comparative  Zoology. 

Bill,  his  family  and  I  saw  much  of  each  other;  I 
became  Secretary-Treasurer  of  the  Boston  Club 
and  joined  the  A.M.U.  It  was  at  that  meeting  in 
Toronto  in  1939  that  I  met  my  husband.  Bill  in- 
troduced us.  In  1941  Bill  drove  his  family  and 
me  to  the  meeting  in  Rockland,  Maine.  After  a 
few  days  there  I  became  engaged  to  H.  B. 
Baker.  We  were  married  in  December,  1941, 
and  I  moved  to  Philadelphia  after  a  honeymoon 
on  Sanibel. 

In  subsequent  years  we  saw  Bill  often  and 
kept  in  close  touch.  He  had  many  friends  and  ad- 
mirers. I  am  happy  to  have  been  one  of  them  for 
over  50  years. 

We  will  never  forget  you,  Bill. 

To  Bill -Cecelia  W.  Abbott 

[Cecelia  W.  Abbott  succeeded  Mrs.  Baker  as 
Business  Manager  of  The  Nautilus,  and  she  often 
accompanied  Dr.  Clench  in  his  later  Itfe  on  his 
collecting  trips  to  Kentucky,  Georgia  and 
Florida  and  more  recent  A.M.U.  meetings.] 

So  much  of  Bill  Clench's  life  was  devoted  to 
helping  new  students  and  encouraging 
amateurs  to  study  and  collect  shells  that  I 
thought  it  only  appropriate  that  a  special  trophy 
be  created  for  shell  shows  that  would  emphasize 
his  keen  interest  and  love  of  land  and  fresh- 
water mollusks. 

The    opportunity    afforded    itself   when    the 


FIG.  1.  Three  proud  winners  of  the  William  J.  Clench  Award 
in  Jacksonville,  Florida,  all  of  whom  created  outstanding  ex- 
hibits on  non-marine  mollusks.  (from  the  left:  Archie  Jones, 
Zida  Kibler  and  Alfredo  Romeu).  Photo  by  William  Kibler. 


Jacksonville  Shell  Club  celebrated  its  25th  anni- 
versary in  1984.  I  lent  a  splendid  sketch  of  Bill 
and  with  the  efforts  and  talents  of  club  member, 
Allan  B.  Walker,  a  handsome  plaque  was 
designed.  Bill's  likeness,  surmounted  by  two  of 
his  favorite  shells -Canthyri a  spinosa  (Lea)  and 
Triop.ns  albolabris  (Say)  graced  the  William  J. 
Clench  Award  "to  the  outstanding  exhibit  of 
land  or  freshwater  mollusks  that  shows  overall 
excellence  and  best  furthers  interest  in  non- 
marine  conchology." 

The  award  has  now  been  given  three  times, 
the  first  one,  in  1984  to  Archie  Jones  of  Miami, 
Florida,  an  old  friend  of  Bill's  and  an  expert  on 
Liguus.  Other  winners,  to  date,  have  been  Zida 
and  William  Kibler,  in  1985,  of  Shallotte,  North 
Carolina,  for  their  worldwide  land  shell  exhibit, 
and  Dr.  and  Mrs.  Alfredo  Romeu  of  Jackson- 
ville, Florida,  for  their  Land  Shell  of  the  World 
exhibit,  1986.  Our  hope  has  been  realized  that 
this  trophy  is  inspiring  others  to  produce 
outstanding,  educational  exhibits,  and  is 
reminding  us  all  of  the  wonderful  qualities  of 
this  kind  and  generous  man  after  whom  the 
Clench  Award  is  named. 


0 


INFORMATION  FOR  SUBSCRIBERS 


The  annual  subscription  rate  for  The  Nautilus 
is  $15.00  for  individuals  (foreign  $18.00)  and 
$20.00  for  institutions  (domestic  or  foreign). 
Subscriptions  may  begin  in  January.  Send  check 
or  money  order  made  out  to  "American  Mala- 
cologists"  to  the  Business  Manager,  P.O.  Box 
2255,  Melbourne,  Florida  32902-2255,  U.S.A. 


Back  issues  from  volume  72  to  date  are  ob- 
tainable from  the  Business  Manager.  Volumes  1 
through  71  (if  available)  may  be  obtained  in 
reprint  or  original  form  from  Kraus  Reprint 
Co.,  Route  100,  Millwood,  New  York  10546. 

Advertising  rates  may  be  obtained  from  the 
Business  Manager  or  Editor. 


CONTRIBUTORS 


Manuscripts:  Authors  are  requested  to  follow 
the  recommendations  of  the  Style  Manual  for 
Biological  Journals,  which  may  be  purchased 
from  the  American  Institute  of  Biological  Sci- 
ences, 1401  Wilson  Boulevard,  Arlington,  Va. 
22209.  Manuscripts  should  be  typewritten  and 
doublespaced;  original  and  one  copy  are  re- 
quired, to  facilitate  reviews.  Tables,  numbered 
in  arable,  should  be  on  separate  pages,  with  the 
title  at  the  top.  Legends  to  photographs  should 
be  typed  on  separate  sheets.  Explanatory  terms 
and  symbols  within  a  dravnng  should  be  neatly 
printed,  or  they  may  be  pencilled  in  on  a  translu- 


cent overlay,  so  that  the  printer  may  set  them  in 
8  pt.  type.  There  is  a  charge  of  50  cents  per 
word  for  this  extra  service.  All  authors  or  their 
institutions  will  be  charged  50  cents  per  line  of 
tabular  material  and  taxonomic  keys.  The  pub- 
lishers reserve  the  right,  seldom  exercised,  to 
charge  $45  per  printed  page. 

An  abstract  should  accompany  each  paper. 

Reprints  are  available  at  cost  to  authors. 
When  proof  is  returned  to  authors,  information 
about  ordering  reprints  will  be  given.  They 
are  obtained  from  Economy  Printing  Co.,  Inc., 
R.D.  3,  Box  169,  Easton,  Maryland  21601-9430. 


MOLLUSK  VOUCHER  SPECIMENS 


It  is  becoming  increasingly  important  for 
future  research  purposes  that  an  identified  sam- 
pling of  species  mentioned  in  publications  be 
deposited  in  a  permanent,  accessible  museum 
specializing  in  mollusks.  This  is  particularly  true 
of  mollusks  used  in  physiological,  medical, 
parasitological,  ecological,  and  experimental 
projects. 


Several  museums  of  natural  history  have  ex- 
tensive modern  facilities  and  equipment  for  the 
housing  and  curating  of  voucher  specimens. 
Material  should  be  accompanied  by  the  identifi- 
cation, locality  data  and  its  bibliographic 
reference.  There  is  no  charge  for  this  perma- 
nent curating  service,  and  catalog  numbers,  if 
desired,  will  be  sent  to  authors  prior  to  publica- 
tion. 


WANTED  -  OLD  SHELL  BOOKS 

Will  pay  good  prices  for  libraries,  second-  Phone  (1-305-725-2260)  or  write:  R.  Tucker 

hand  books  and  reprints  on  mollusks,  shells  Abbott,  American  Malacologists,  Inc.,  P.O. 

and    conchology.    Back    numbers    of    The  Box  2255,  Melbourne,  FL  32902.  Free  ap- 

Nautilus,  vols.  40-71  wanted,  $1.50  each.  praisals. 


americahmalacologists.  mc. 

PUBLISHERS  OF  DISTINCTIVB  BOOKS  ONMOLLUSKS 

THE  NAUTILUS  (Quarterly) 

MONOGRAPHS  OF  MARINE  MOLLUSCA 

STANDARD  CATALOG  OF  SHELLS 

INDEXES  TO  THE  NAUTILUS 

(Geographical,  vols  1-90;  Scientific  Names,  vols  61-90) 

REGISTER  OF  AMERICAN  MALACOLOGISTS 


MBL  WHOI   LIBRARY 


UH    17YE    7